it Bath Dap cTeake Ud Meth ct f (8 al bh Ale 4 Hayy, chad Weed CTReEGS Ieee Rat We atc aa yi HAWS 5 este La Soe Ghia te ey oe) y _ 2 Res * : Ay tie 7 Mee tS i) aa aan . PROCEEDINGS ~ BIOLOGICAL SOCIETY ala | _ WASHINGTON VOLUME 111 NUMBER 3 18 SEPTEMBER 1998 ISSN 0006-324X THE BIOLOGICAL SOCIETY OF WASHINGTON 1998-1999 Officers President: Richard P. Vari Secretary: Carole C. Baldwin President-elect: Brian F. Kensley Treasurer: T. Chad Walter Elected Council Michael D. Carleton Rafael Lemaitre W. Duane Hope Roy W. McDiarmid Susan L. Jewett James N. Norris Custodian of Publications: Storrs L. Olson PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: Frederick M. Bayer Invertebrates: Stephen L. Gardiner Plants: David B. Lellinger Frank D. Ferrari Insects: Wayne N. Mathis Rafael Lemaitre Vertebrates: Gary R. Graves Membership in the Society is open to anyone who wishes to join. There are no prerequisites. 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Payment for membership is accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards. ; Manuscripts, corrected proofs, and editorial questions should be sent to: EDITOR BIOLOGICAL SOCIETY OF WASHINGTON NATIONAL MUSEUM OF NATURAL HISTORY WASHINGTON, D.C. 20560, U.S.A. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Periodicals postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. © This paper meets the requirements of ANSI/NISO 239.48-1992 (Permanence of Paper). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):473—-503. 1998. Horton H. Hobbs, Jr. (29 March 1914—22 March 1994). Biographical notes Karen Reed and Raymond B. Manning Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. Abstract.—The 211 papers and abstracts published by the late Horton H. Hobbs, Jr., are cited and annotated by a list of new taxa published in each paper. An alphabetical list of taxa named by Hobbs is provided; the repository and the catalogue number are cited for each holotype. Hobbs’s reminiscences of his introduction to crayfishes and his early work are summarized from an oral interview. To many who knew him and his work, the name Horton H. Hobbs, Jr., (Figs. 1, 2) is synonymous with freshwater crayfishes of North America. Indeed, to many he was known affectionately as “‘crawdaddy.”’ The enormous advances in our understanding of the systematics, distribution, and evolution of crayfishes that resulted from his studies in a career spanning six decades are well known to all astacologists. Perhaps less known are his contributions on other groups, including freshwater and cave shrimps and crabs, and ostracod associates of crayfishes. Hobbs’s studies resulted in the recogni- tion of many new taxa: | new family (Cam- baridae); 38 new genera and subgenera (1 genus of palaemonid shrimp, Neopalae- mon; 8 genera of entocytherid ostracods; and 29 genera and subgenera of North American crayfishes); and 286 species (168 crayfishes, 104 entocytherids, 8 caridean shrimps, and 6 freshwater crabs). A remarkable number of Hobbs’s contri- butions are monographs or other book- length works on a wide variety of topics: 1942d, crayfishes of Florida, 179 pp.; 1964a, Hobbs & Villalobos, crayfishes of Cuba, 59 pp.; 1969a, Chace & Hobbs, freshwater and terrestrial decapods of the West Indies, 258 PPp.; 1971b, entocytherid ostracods of Mexico and Cuba, 55 pp.; 1972a, subgenera of Procambarus, 22 PPp.; 1972g, crayfishes of North and Middle America, 173 pp.; 1974c, checklist of North and Middle American crayfishes, 161 pp.; 1977b, Hobbs & Peters, entocytherid os- tracods of North Carolina, 73 pp.; 1977c, Hobbs, Hobbs III, & Daniel, trog- lobitic decapods of the Americas, 183 pp.; 1981b, crayfishes of Georgia, 549 pp.; ~ 1982e, Hobbs & Hart, revision of Atya, 143 pp.; 1986a, Andolshek & Hobbs, entocytherid ostracods of Georgia, 43 pp.; 1987b, revision of Astacoides, 50 pp.; 1989g¢, illustrated checklist of American crayfishes, 236 pp. Hobbs’s productivity led a European stu- dent to remark to one of us (R.B.M.) “‘Oh, Hobbs. He writes faster than I can read.”’ Hobbs III recalls (in litt.) that during the late 1930s, Hobbs’s father held him by his ankles while Hobbs slithered into a vertical crack (River Bed Cave, Columbia County, Florida) to collect Procambarus (O.) palli- dus—he then pulled him back out (for other anecdotes by and about Hobbs, see Hobbs 1986d). 474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 7° a Fig. 1. Hobbs collecting after the 1967 meeting of the Association of Southeastern Biologists, Columbia, South Carolina (J. F Fitzpatrick, Jr. and Hobbs III). J. F Fitzpatrick, Jr. (in litt.), provided two route on some field excursion. Because he other anecdotes. ““One day, in the ‘60s, I was in motion, he did not call or write think—Hobbs had stopped by to see Miss ahead to announce his arrival. When he Peggy on some business while he was en knocked on the door in the middle of the VOLUME 111, NUMBER 3 day in an active residential neighborhood of a small Virginia city, he found Miss Peggy home alone. The entire conversation, not short, was conducted with him standing on the porch and she behind the cracked door with the chain engaged. To her, ‘It just wouldn’t be proper to have a man in the house with Miss Lucille away on an er- rand’, even such a true gentleman of long acquaintance as was Hobbs. He delighted in relating this story to those who knew her as an example of her propriety, but he never faulted her behavior; he only respected it in his amusement.” “The other goes back to his graduate stu- dent days at Gainesville and reveals a side of his character that only those of us who knew him well knew. He and Archie Carr were classmates and shared an office with a few others. Despite the reticence of his later years, a youthful Archie apparently was animated and outspoken. He had a hab- it during repeated discussions of rearing back on the hind legs of his wooden chair, and when he made an emphatic point, he would slam the front legs on the floor. The mischievous crew connived to play a trick on Archie with Hobbs surely one of the conspirators if not the actual perpetrator. They got a couple of fireworks ‘torpedoes’ and clandestinely attached them to the front legs of Archie’s chair. When he came in, they led him into a topic to which they knew he would react strongly. True to form, he reared his chair and at an appropriate point slammed the front legs. Archie was attached to the ceiling by his nails as the others rolled on the floor in merriment. De- spite this treatment, the group remained friends for their whole lives.” All of us who knew and worked with Horton have many fond remembrances of him. He was the quintessential southern gentleman, always rising when a woman entered his presence. It pained him not to be able to open doors for women, and his female car-poolers were trained to wait at a door if he fell behind them. Elizabeth Nel- son remembers that in his last years, in frail 475 condition and walking precariously with a cane, he rushed to open a door for her as she walked by with a small package. She knew that to refuse his offer of assistance would have pained him terribly. Elizabeth also remembered his expres- sion, a mixture of embarrassment and delight, when he was presented with a pair of boxer shorts with two flies at his retire- ment party—the idea being that having studied entocytherid ostracods so long he might have developed hemipenes. Brian Kensley joined Horton on one of his last field trips. “In the fall of 1987, I accompanied Horton on a field trip to east- ern Texas. He wanted to complete a survey of the crayfishes of this area, and also gath- er some data on Fallicambarus devastator Hobbs & Whiteman, 1987c, the burrowing crayfish that causes much destruction to fields and lawns. We drove in Horton’s car. He would set the speed control, and dis- course on the flora and geology of the area Wwe were driving through. Every now and then we would have to stop, so that he could push a net through a puddle next to the road, or to show me a tree with which I was unfamiliar. Between whiles, he re- counted incidents from his early days of field collecting and teaching. It was for me a natural history revelation. Arrived in Tex- as, we met with farmers and agricultural ex- tension officers to gather information on devastator, and to dig in the bone-dry, con- crete-hard fields dotted with chimneys. This was my primary role. Once I had got down to the water table, Horton would lie stretched out on the ground, one arm reach- ing 2—3 feet down into the hole for the cou- ple of crayfish at the bottom. We also did a lot of (much easier) digging in stream banks, and this is where I uncovered spec- imens of a beautiful spotted crayfish which he eventually named for me [Procambarus (Girardiella) kensleyi Hobbs, 1990a]. Hunt- ing season opened while we were in the Neches River area. On the third occasion while we were either walking through the woods, or working at a stream bank, and a 476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 111, NUMBER 3 rifle shot rang out nearby, Horton turned to me (I was distinctly edgy at this point) and said: “‘Let’s go home’, which we did, all the way back to northern Virginia, at the sedate speed limit while I absorbed yet more land- scape science.” Horton thoroughly enjoyed his martini at cocktail hour (see Fig. 3, showing Horton on his boat, the martini barge, on Lake Bar- croft). While one of us (R.B.M.) was living in Tunisia, Horton graciously shared his home with me. On one of my trips back, I noticed that Horton was having a clear cocktail in a 12 oz. glass rather than in his normal martini glass. This appeared strange to me, but I didn’t comment on the large drink, assuming that Horton’s affinity for martinis had expanded exponentially. Someone told him of my concern, and he had many laughs from the occasion, as he had shifted from martinis to a gin and tonic, with lots of ice, for the summer. I suspect that I took Horton his last martini during his last stay in the hospital. It was in a glass jar, which he gleefully accepted and hid in a drawer so that he could have it at happy hour. Later that day, his daughter Nina vis- ited him, and Horton told her “‘look what I have.”’ She smelled the jar and asked where he got it, probably knowing full well that I was involved. Aspects of Hobbs’s life and career have been summarized in other obituaries by Fitzpatrick (1995a, 1995b, 1996) and Hoff- man (1994). Hobbs’s southern manners were characteristic of him. One aspect of his life that was known to those who shared dinner as his guest was his interest (and ability) in baking and cooking. Alan David- son (1979:432) in North Atlantic Seafood published his recipe for hush puppies. Da- < Fig. 2. 477 vidson referred to him as “The scholarly Mr Hobbs.” Cooper & Cooper (1997:616) acknowl- edged Hobbs’s influence: ‘“‘As always, we are immensely grateful to the late Horton H. Hobbs, Jr., for the splendid lessons he taught (not all of them about decapods).”’ Only his colleagues in the Department of Invertebrate Zoology at the National Museum of Natural History are aware of one highlight of Hobbs’s career. In 1976 Thomas E. Bowman and Louis S. Kor- nicker, members of the department’s self- appointed SOL Awards Committee, awarded him the “‘Smithsonian Order of the Lobster.’’ The award celebrates an ac- complishment or achievement of outstand- ing insignificance, something that usually is quite difficult to associate with Hobbs. The award was based on the following se- ries of misadventures. Hobbs and colleagues C. W. Hart, Jr. and Margaret Walton introduced four new names for the entocytherid ostracod, Don- naldsoncythere donnaldsonnensis (Klie, 1931) (see Hobbs & Peters 1977:43, 44 for a synonymy), already burdened by two synonyms, Entocythere humesi Hoff, 1943 and E. pennsylvanica Hart, 1960, as well as four unavailable names published as erroneous spellings: E. donnaldsoni Wolf, 1934-1938; E. donalsonensis Rioja, 1943; and E. donaldsonensis Tressler, 1947. Hobbs & Walton (1961a) named E. hiwasseensis from Georgia; Hart & Hobbs (1961b) named E. tuberosa from Tennes- see (with the erroneous spelling tubercu- lata in the same paper); Hobbs & Walton (1963b) named Donnaldsoncythere scalis from Virginia and D. ileata from Virginia. United States crawfish workers, meeting of Association of Southeastern Biologists, Memphis, Ten- nessee, 19 April 1969 (includes all significant workers with crayfishes and their associated biota, except for Perry C. Holt, who worked on branchiobdellids). From left to right, C. W. Hart, Jr., J. E Fitzpatrick, Jr., Glen Gentry, Rudolph Prins, Daniel J. Peters, James E Payne, Raymond W. Bouchard, Jean E. Pugh, Horton H. Hobbs III, Horton H. Hobbs, Jr, Marilyn Black, Martha Reiser Cooper, Joe B. Black, and John E. Cooper (J. E Fitzpatrick, Jr., R. Prins, and Hobbs III). THE BIOLOGICAL SOCIETY OF WASHINGTON S OF . J SEDINC PROCEI 478 polepun ‘ ‘(sisjag “f [atueq) winisjog ‘sroqnq uear Aq ojoyd ‘Og6] Aine p ‘Pley oy) ur ‘(siojog “f JorURC) Payepun ‘SuruuLPY YIM d31eq IUNAeU Oy) UO ‘(T]] SqqoH) PIP oy) Ur ‘(TIT SQqOH) parepun ‘sKep aj[tAsourey ‘(IIT SqqOH) 6961 ‘YUsyAvio v SurAolugq sya] Joddn woy ‘asimyoo[D If ‘sqqoH “H UOMOR, “¢ “BI 3 = A TAP 2 So Pe i a a, VOLUME 111, NUMBER 3 Biographical Notes Here we reproduce part of an Oral His- tory Interview of Hobbs by Pamela M. Henson, Smithsonian Institution Archives, in July 1976. It and the rest of the inter- view, largely dealing with aspects of his ca- reer after he left the University of Florida to go to the University of Virginia in 1946, form Record Unit 9509 in the Smithsonian Institution Archives. The questions are by Henson, the answers by Hobbs. Question: As we discussed, we can start with some biographical information about you and your education, and how you came to be interested in Crustacea. Answer: My undergraduate, well, all of my work was done at the University of Florida. It was during the depression years, so when I received my Bachelor’s degree I was able to get an assistantship in the de- partment and took my Master’s. There was no place to go, so I stayed on and joined the faculty after I took my Master’s, teach- ing the introductory courses in biology. I stayed on for my doctorate; and it took me several years because of residence require- ments and working full time, so I got my degree in 1940. I spent one summer at the Mountain Lake Biological Station during that time, doing work with Dr. [Chauncy M.] Gilbert, a course in arthropods. You asked about my interest in crayfish- es. This began when I was a freshman in college. We came to the laboratory exercise dealing with the crayfish; the first day it was external anatomy—had no difficulty whatsoever, everything was beautiful. The second day had to do with internal struc- tures, and the crayfish that I was given ap- parently had demised with the flood, be- cause it was complete soup inside. So I sup- pose I was an eager beaver, in part, and I knew that not far from the boarding house where I was living at the time there was a little creek. I went down after school that afternoon and I caught a few crayfishes and brought them back to the room. That eve- ning I made my dissection and everything 479 was as it should be, but I had some crayfish left over and I couldn’t see killing them. So I found a jar to put them in (I planned to take them back to the creek the next eve- ning) and put them on my desk where I was studying. In a few minutes I noticed a com- motion, and a male and female had gotten together. I watched them for a while, and I said, ‘““Well, maybe I’ll get eggs in a few days.’ And sure enough, within two weeks, eggs were there. About three weeks later the female was carrying the young, and the more I watched them the more interested I became. By the end of that semester, I went to the chairman of the department and told him that I’d become interested in crayfishes and would like to do some work with them. He said, “‘Well, I know nothing about them but I’ll be happy to help you in any way that I can.”’ So that’s the way it began, and since 1931, I’ve looked at not much else. Question: That is interesting. And I sup- pose your room turned into a big aquarium? Answer: No, not really. But strangely enough, I was never able after that time to rear a clutch of eggs to adulthood until after I came to the Smithsonian. Question: Just chance? Answer: I didn’t know enough about them at that time to do the wrong thing! Question: That’s amazing. But there was no one at the university studying crayfish? Answer: No, no, there was only one per- son in the United States at the time. The big crayfish men had died: [Walter] Faxon and [Arnold Edward] Ortmann; and Edwin P. Creaser at Michigan Museum [Museum of Zoology, University of Michigan] was the only person working. That brings me to my first visit to the Smithsonian [Institution] which might be of some interest to you. In 1935, I went up to the Mountain Lake Biological Station for the summer. I had a letter of introduction from my major professor [J. Speed Rogers] at the University of Florida, and one from Dr. Ivey E Lewis, who was Dean of the University of Virginia, introducing me to Dr. Waldo [L.] Schmitt [then Head Curator 480 of Zoology at the U. S. National Museum]. So after my stay at the station, I came up to Washington hoping that I might be able to examine the crayfishes. Dr. Schmitt took one look at me ... I know you wouldn’t believe it now, at one time I looked a little young. At the ripe old age of twenty-one I suppose I looked like I was about fifteen. He told me that he would be glad to have me look at the crayfishes, but that I could not open any of the types, couldn’t open the bottles. Well I had made that trip at some sacrifice, and Dr. Creaser couldn’t identify my material. Dr. Schmitt was very gracious but not of much help in doing so. Not until 1937, when I convinced Dr. Schmitt that I was a serious worker, did he allow me to come back and examine the type specimens. So I worked for five years on crayfishes not knowing the name of a single crayfish that I was working with! Question: Oh my goodness! Yes, not having compared them with the ones here. Answer: I needed to compare them with the types. Question: Well, you must have come up with some original observations. Answer: Oh, well, a number of my ani- mals were new. When I started working in Florida I think there’d been about, oh may- be, four species recorded from the state; and when I finished there were forty-two of them. Question: [Laughter] That’s a difference! Answer: Which labels me perhaps as a splitter. Question: Yes, but still, there just hadn’t been that much work done. There was a paper written by Fenner [A.] Chace [Jr.] at one point about the lack of work in system- atics being done in that area during that pe- riod [reference to Chace 19517]. I guess it was not one of the more worked on areas? Answer: No, it never has been. Crayfish- es have never attracted very many people. Those who start, most of them have fallen [by] the wayside after a little bit of work. I’ve worked on crayfishes I suppose longer PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON than any other human ever has, in terms of number of years spent. . Question: What was the Smithsonian like when you first came here to work on those? Answer: Oh, it was a delightful place. Of course, my coming from a small town and coming to Washington, in part, I suppose I was much impressed. Dr. Schmitt was most gracious, as were Mr. [Clarence R.] Shoe- maker, Mr. [James O.] Maloney, and Miss [Mary Jane] Rathbun. Question: She was still here? Answer: She was still here. I can’t re- member whether I met her or not, but at least I saw her. And Dr. [Leonhard] Stejne- ger was a marvel. All of them were most gracious and helpful, and as I said, I do not hold it against Dr. Schmitt at all because I suppose I did look like a high school kid who thought he was going to do something. Question: Right, take apart all the type specimens. Answer: With this particular group, the group that’s dominant in Florida, unless you look at the first pleopods of the males you can’t identify the crayfish at all, and through a bottle you couldn’t see them. Question: Were the collections of cray- fish at that point fairly extensive here? Answer: Well certainly nothing in com- parison with what we have at the present time. It was perhaps not even the most im- portant collection in the country. The Mu- seum of Comparative Zoology had many more types than we had at that time; and there were a few at the Philadelphia Acad- emy [of Natural Sciences], but a very im- portant collection existed here, one that formed a nucleus around which we’ ve been building on for a number of years. Now, of course, we have the largest collection of crayfishes in the world—perhaps larger than all the other collections of crayfishes in the world combined [the collection of crayfishes now includes about 25,000 lots and 1500 lots of types]. Question: Was anyone working on them then? Answer: No, no one at the Smithsonian VOLUME 111, NUMBER 3 had worked on crayfishes. I’m the first per- son. Question: Yes, to come in and work on it. Answer: Faxon worked at the Museum of Comparative Zoology, and Ortmann was at the Carnegie Museum, and Creaser was at the Michigan Museum. Perhaps you know that we have the Michigan collection now. After I came here my major professor be- came director of the Michigan Museum, and he said, ““Nobody’s working on cray- fishes out here, and we’re crowded. Wouldn’t the Smithsonian like to have it?” That was just before I came here; I was in Charlottesville at the time. He said, ““You’re nearby so you can use them from time to time.”’ So we inherited that collec- tion. Question: How easy was it to start work- ing in a field like that where no one else was working? Answer: As I look back on it, I had a delightful time all the way through. I knew that I could tell the difference between what I had, and just because I didn’t have a name for them at the time didn’t bother me too much. The literature had not been cluttered with my group of animals that I was work- ing on in Florida, so there was very little in the literature at all. So there was no great literature problem to face. Question: Yes, going through the masses of descriptions. Answer: I hate to say what has been done to it since. [Laughter.] As I frequently say, they will curse me and say all kinds of things, but they can’t ignore me any longer with the crayfish. Question: Right, you have left your mark. Were there avenues, let’s say, for publishing your results? Answer: Oh, they were very limited, of course. Again, I came along during the de- pression years, but Dr. Schmitt was very, very kind, and I can tell you one lovely little anecdote that happened to me in con- nection with publications. I used the Pro- ceedings of the Florida Academy of Sci- 481 ences—it used to be called the Journal. I used that for some of my work. Then the Charleston Museum at that time was pub- lishing a series of Leaflets and Zoologica out of New York, and The American Mid- land Naturalist [also were available]. So there were a number of places that were open. But I sent one of my early papers up to Dr. Schmitt, describing seven new cray- fishes from Florida—lI think it was seven new ones—and redescribing, in essence, a species that had already been described. In this I used the word “‘crawfish.”’ I'd always used crawfish as a name, and throughout most of the South they’re still referred to as crawfish. Dr. Schmitt carefully went through my manuscript completely and changed it to crayfish everywhere. So I de- cided that, well, if they’re going to force me to do this (I didn’t raise any questions whatsoever), Ill just accept it. So since that time I have used “crayfish,” and most of my students do. However, most of the other people, particularly those who’ ve worked in the South, still write crawfishes when they write it. Question: 1 hadn’t even realized that there was the difference in words. Answer: Oh yes. And one other thing oc- curred in that paper. I had caught crayfishes from a cave from which they had been re- ported in Florida. Faxon had identified the animal as Cambarus lucifugus [error for acherontis sensu Faxon, not Lonnberg; re- placed by lucifugus Hobbs (1940a)], a spe- cies that had been described by [Einar] Lonnberg in the latter part of the nineteenth century from a well down near Orlando. Well, believing that the written word was infallible, I assumed that the specimens from this cave were what Faxon said they were. This was in the same manuscript, and Dr. Schmitt passed it on up to Dr. Stejneger, who at that time was Head Curator. Dr. Stejneger took one look at it and he said, ‘“What right does he have to say that this is lucifugus?” [error for acherontis sensu Faxon.] He said, ““He should go down to that well and catch some crayfish from that 482 place to be sure.”” So Dr. Schmitt wrote back to me and gave me Dr. Stejneger’s re- marks. Well I was a little bit upset because I thought the possibility of my finding that well that had been dug back in the latter part of the last century was very slight, and Florida was not easy to get around in, it was certainly not like it is now—all highways. But a friend of mine, Lewis J. Marchand, who lived down near Orlando, happened to come by my office within a day or so after I'd heard from Dr. Schmitt. I asked him if he had ever seen any white crayfishes down in that area. He said, ““Oh sure, I know a spring where I’ve seen them a number of times. So I said, “How soon can you be ready to go?’’ And he said, “‘Well, tomor- row morning will be fine.’ So I said, “Well, we’ll start then.’’ Palm Springs is what it was. In the meantime, I had another friend, fellow student, at the university who was very good at water goggling—that’s what we called it in those days—it’s mod- ified scuba diving but you don’t have air and so on. Anyway, I asked him if he didn’t want to go. This was a cold November day, and, believe you me, Florida can get cold during the winter, the northern part. So we went down to Palm Springs, and when we got to this little spring I looked down—of course it hadn’t been used since summer— it was covered with algae, and lying on the algae were white crayfish everywhere. So this friend of mine who went along with us took his gear and jumped off into the spring and time after time he came up, so we got forty-four of those animals that day. I got back to the laboratory, and sure enough, it was not the same as the thing from the cave, and was precisely what Lonnberg had described. So I had to redescribe it and put a new name [Cambarus lucifugus]| on the material from the cave. Question: Yes. You can’t ever count on Answer: So Dr. Stejneger sitting in Wash- ington certainly saved me considerable em- barrassment, and taught me a good lesson not to accept the printed word. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Question: That is interesting, and you were having things published. Did you start collecting yourself at that point? Answer: Oh, I started collecting back in 1931, when I was still a freshman. Question: You did keep track of your dif- ferent types? Answer: Oh yes. One of the biggest helps, I suppose, was Dr. [J. Speed] Rogers, my major professor and the chairman of the department at that time, with his meticu- lousness in keeping notes and insisting that everything be carefully curated. So I started out being trained as a curator, I suppose. Through the years I amassed a collection of some 80,000 specimens that I brought with me when I came to the Smithsonian. All of those were catalogued and we’re still using my old numbers. The collection’s so tre- mendous that we haven’t been able to in- corporate nearly all of them into the Smith- sonian catalogue, so we’re still using my old cards. Question: Where did you keep it all? Answer: In my basement at home. Question: You did? Answer: I had a tiny little office at the University of Virginia in an old building. The office was about the size of this little anteroom out here. In one of the rooms the floor fell in, the basement where I had put so many crayfish. I had to take everything out and have the floor reconstructed to sup- port the crayfish collection. Question: Yes, I guess everything was pretty much alcoholic storage? Answer: Everything was alcoholic stor- age, yes. Question: Which is fairly heavy. The professors you were working under were they systematics people? Answer: All of them were systematists, for the most part. There were four men in the department at the time, four of the full professors. All of them were graduates of the University of Michigan. One of them had taken his doctorate at, I believe, New York—I’m not sure whether it was Colum- bia [University] or not—but one of them VOLUME 111, NUMBER 3 had taken his doctorate elsewhere. But all of them had been trained in Michigan under [Alexander G.] Ruthven and [Robert W.] Hegner. Three of them were entomologists: Dr. Rogers worked on the Tipulidae, the crane flies; and Dr. [Theodore Huntington] Hubbell on the Orthoptera, grasshoppers, and Dr. [C. Francis] Byers on the Odonata, dragonflies. The fourth member was a mammalogist. He and I were good cronies because he was interested in bats—primar- ily, and bats live in caves and crayfish live in caves. So we had a delightful time team- ing up going on field trips. I'll tell you one other story of Dr. [Harley Bakwel] Sherman, the mammalogist. The library at the University of Florida had got- ten a new photostat machine and had dis- carded the old one. Dr. Sherman and I res- cued it and rebuilt it. We had heard that there had recently been aerial photographs taken of the area around Gainesville, of the entire county, that were available in the county agent’s office. So we borrowed these, and made copies and placed them in our notebooks. Fortunately, those photo- graphs had been taken during the winter months when the deciduous trees had lost their leaves. This meant that where you would see a black spot on our maps this was a Cluster of live oak trees, and live oaks usually grow along some depression, ‘fre- quently indicating a sinkhole or maybe a cave. In Florida, that section of the state is quite flat and the roads are built on the sec- tion lines so that they run at mile intervals, almost straight, occasionally going around a sinkhole or something of the sort. But no place are you more than a half a mile from any spot—if you were on the road—from any place within the quadrangle. So we would ride down the section lines with our maps, and if we’d see a black spot, we'd get out of the car and look to see whether this was a cave in which there might be bats or crayfishes or something of the sort. One afternoon we’d been out, it was get- ting quite late and there were two graduate students with us, one of them, Dr. [Jerome] 483 Krivanek, who’s now at Vanderbilt [Uni- versity], and a young man, [William M.] McLane, who has recently died. Dr. Sher- man was driving, and I said, ““Dr. Sherman, here’s a little place right close to the road. Stop, it won’t take Billy and me a moment or two to have a look.”’ So we rushed over. It was nothing but a depression, but on our way back to the car there was a perfectly cylindrical chimney that went right down, dropped down, oh, between fifty and sev- enty feet. I peeked over the edge of it and saw that there was a little water in the bot- tom of it which excited me, so we rushed back to the car and asked them if we didn’t have time to make one quick drop into this hole to see what it was. McLane and I went down into the hole and the other two stayed up above. We got down to the bottom, and I saw white crayfish on the bottom. We looked around and there was a little open- ing into the side, about two and a half feet in diameter; and we crawled into this open- ing, and that led into a fissure that was about four feet wide and some sixty or sev- enty feet long, with no floor, but the entire thing with water under us. It was shallow at one end, then it dropped off rather quick- ly. Even to this day we don’t know how deep it really goes, it just continues on. I saw the white crayfish down there and I turned to this graduate student, and I said, “Billy, if you’ll catch one of those cray- fishes and it’s a new one, Ill name it for you.” Well, I knew what they were so I was perfectly safe. So Billy jumped into the wa- ter with a dip net and started scurrying around, but he was missing them. I said, ‘““Well, let’s go.’ The water had gotten so cloudy you couldn’t do anything. So we crawled back up to the top, and when we reached the top of the cave, he turned to me and said, ‘““Well, Doc, I didn’t get any of the big ones but I got this little one.”’ He handed me a vial with a little tiny crawfish [Troglocambarus maclanei; in his dedica- tion of this species to McLane, Hobbs (1942b:349, footnote) commented: “It is a pleasure to name this species for Mr. 484 McLane, who has been a companion on many collecting trips, and who has added numerous valuable specimens to my collec- tion.’’] in it, the most amazing animal I had ever seen in my life. I couldn’t believe that it was anything except a mutant or some- thing had gone wrong somewhere. We went on back into Gainesville, and I immediately got the binocular scope on it and saw that it was really something out of this world. So the next day I was anxious to get back and get some more of them. Most of the young men who usually went with me were tied up in classes, they couldn’t go. I found one man who’d go, and we had gone out to the car; we had our ropes and collecting equipment and so on. About that time I spotted the same man who had caught the crayfish down in Palm Springs, going across the campus. I called to him, and told him that we were going out to a cave, and wouldn’t he like to go, and he said no, that he had an organic exam the next day. ““Well, that’s too bad,”’ I said, “it’s a wonderful place to water goggle.” I knew that would get him, no question. “Oh?,”’ he said, “All right, Pll go.”’ I called him Jelly, so that gives you some idea as to his size. He wasn’t all that obese, he was pretty wide, he had some flesh on him and I was worried all this time about his getting through that side of the pit. Question: That’s true, yes. Answer: But, we got out to the cave, we went down it, . and he said, ““Now where’s that place to dive?’ I said, “It’s right through there.’’ He looked at the hole and he said, “I can make it.’’ So into the hole he went, all of us. It was such that we could prop our feet against one wall and lean our back against the other one, no place to stand whatsoever. He said, ‘““Now where’s that place to dive?”’ I said, ““You’re there.”’ Well, if looks could have killed any- one, well of course, I would have been dead. But he was a good sport so he said, ‘All right.”” So he put on his goggles, and he went down, and he came up time after time with this big white crayfish, Procam- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON barus pallidus, which I had recognized all along—but none of the little fellow. He said, ““Well, I’ve about had it.’’ I said, “‘Jel- ly, go down just one more time,”’ and this time when he came up he rotated just be- fore he surfaced and his light beam hit the submerged ceiling, and he saw one of them and picked it off and brought it up. After that, he went down and he got either three or four more that afternoon, all of them col- lected from the ceiling. So this little animal was highly adapted for living on submerged ceilings of caves; down below it would have been in competition, with a much less chance of survival certainly than it has above, a very small animal. What had hap- pened, of course, McLane, when he jumped into the water, had jarred the water so that the animal became dislodged and on the way down Billy happened to catch it. Question: Going down without ever looking up. . Answer: I finally caught one when Mar- chand, the man who had caught so many of them, jumped in one day and another one was dislodged and I saw it come loose and grabbed it with my dip net. That’s the only one I’ve ever caught. Question: But it was, I guess, a com- pletely new type? Answer: Oh, it was a completely new ge- nus [Troglocambarus]. It is the most fan- tastic crayfish that we know at the present time. Most crayfishes have teeth on their third maxilliped for chewing. In the first place the maxillipeds have tremendously large and long setae that interlock, and by carrying water through the gill chamber over this setal net, it filters its food out of the water. There is no other crayfish that utilizes this technique. Question: Fascinating, too, that you did find it. Were there many students in system- atics at Florida at that time? Answer: At that time, ecology was the thing, as it has become in recent years. I was trained as an ecologist; of course, the kind of ecology that we did then had little resemblance to the kind that exists at the VOLUME 111, NUMBER 3 present time. All of us were a combination systematist and ecologist. I was trained in ecology, and the only reason I got into sys- tematics was that I couldn’t identify my an- imals and there was no one to help me, and I had it to do. Question: Yes, because that had not been done before. So you stayed there until 1946? Answer: That’s right. Question: And then you moved to the University of Virginia. Answer: While I was there I came up here many weekends to work in the collec- tion. Question: You were using these collec- tions? Answer: Oh, yes, I was using them from the outset. At one time Dr. Schmitt was able to get a little funds to help me to come up and work on weekends. At that time, too, he did another very nice thing for me. Most of the people had to be out of the building by a certain hour, and I was permitted to stay on until midnight. Some way he man- aged to help me out to that extent, because it was so rare—I could only work on Sat- urday night. Sometimes I would come up on Friday, I could work Friday night and Saturday night, or on holidays when I came. Question: I guess you got to know the collections here fairly well. Were they in fairly good condition—identified or cata- logued? Answer: Oh, yes, all of the old material was, and much of the time that I was here, when I would come up on weekends, I was working up collections that had accumulat- ed and identifying them so that they could be catalogued by the time that I came back and they were very nice in sending me du- plicate cards for everything that I identified. Question: Were there many collections coming in during those years, were there expeditions or collecting trips? Answer: No, not a great many. There had been a considerable backlog because, as I said, no crayfish man had been here since [William Perry] Hay worked at the muse- 485 um. He also taught high school here in the Washington area, as I understand it. He’s one of the few older members—crayfish people—that I met, but he had retired and was quite an old man when I met him. He lived in Florida and made a special trip to come up to Gainesville to see me one time, so I was delighted. He subsequently gave me his library, so I have many of his old notes, things that will go to the archives eventually. Co-authors of Horton H. Hobbs, Jr. Hobbs published many papers with co- authors, acknowledging help in the field and collaborating with students and col- leagues. Here we list his co-authors and, where possible, their institution (usually as of the dates), to demonstrate the breadth of his association with others with similar in- terests. As pointed out by Hoffman (1994: 37) some of his papers are co-authored “sometimes for no other reason than to re- ward the collector of a new species.” Andolshek, M. D. (1986a). Hobbs’s research assistant at the museum. National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. (see also Margaret A. Daniel). Banner, Albert H. (1959d). Hawaii Institute of Marine Biology, University of Hawaii, Kaneohe. Barr, Thomas C., Jr. (1960b, 1972b). University of Kentucky, Lexington. Bedinger, M. S. (1964b, 1965a). U.S. Geological Sur- vey. Bouchard, Raymond W. (1973a, 1976b, 1994). Acad- emy of Natural Sciences of Philadelphia. Brown, Arthur V. (1987d). University of Arkansas, Fayetteville. Burr, Brooks M. (1984b). Southern Illinois University, Carbondale. Carlson, Paul H. (1983c, 1985a). Department of Health and Environmental Control, Columbia, South Car- olina. Chace, Fenner A., Jr. (1959d, 1969a). National Mu- seum of Natural History, Smithsonian Institution, Washington, D.C. Cooper, Martha R. (1972e, 1980c). North Carolina State Museum of Natural History, Raleigh. Daniel, Margaret A. (1977c) (nee Margaret D. Andol- shek). Fitzpatrick, Joseph EF, Jr. (1962d, 1970a, 1971d). Uni- versity of South Alabama, Mobile. Hobbs’s last stu- dent. 486 Franz, Richard (1983a, 1986b, 1991b, 1992). Florida State Museum, University of Florida, Gainesville. Freeman, Harry W. (1956d). College of Charleston, South Carolina. Grubbs, Andrew G. (1982b, 1986c). Southwest Texas State University. Hall, Edward T., Jr. (1969d, 1972f, 1974d). Georgia Water Quality Control Board, Atlanta. Hart, C. Willard, Jr. (1956e, 1959b, 1961b, 1966d, 1982e). National Museum of Natural History, Smithsonian Institution, Washington, D.C. One of Hobbs’s students. Hobbs, Horton H., III (1962a, 1970c, 1973c, 1976e, 1977c, 1989e, 1990b, 1991d, 1995a, 1995b). Hobbs’s son, referred to herein as Hobbs III. Wit- tenberg University, Springfield, Ohio. Holt, Perry C. (1967d, 1968b). Virginia Polytechnic Institute, Blacksburg. Hobbs’s first student. Fitzpat- rick recalls that Holt liked to refer to himself and Fitzpatrick as “alpha and omega.” Hubricht, Leslie (1959d). Missouri Botanical Garden, St. Louis; Louisville, Kentucky. Lee, David S. (1976c). North Carolina State Museum of Natural History, Raleigh. Mackin, J. G. (1959d). Texas A&M University, Col- lege Station. Manning, Raymond B. (1977d). National Museum of Natural History, Smithsonian Institution, Washing- ton, D.C. Marchand, Lewis J. (1943a). University of Florida, Gainesville. Massmann, William H. (1952b). Virginia Fisheries Laboratory. McClure, Auden C. (1983e). McLean, Virginia. Means, D. Bruce (1972c). Tall Timbers Research Sta- tion, Tallahassee, Florida. Page, Charles H. (1953b). Charlottesville, Virginia. Parish, Claude E. (1949b). University of Alabama. Penn, George Henry, Jr. (1958h). Tulane University, New Orleans, Louisiana. Perkins, E O. (1967c). Virginia Institute of Marine Sci- ence, Gloucester Point. Peters, Daniel J. (1977b, 1979a, 1982c, 1989b, 1991c, 1993). New Horizons Governor’s School for Science and Technology, Hampton, Virginia. Pflieger, William L. (1988c). Fish and Wildlife Re- search Center, Missouri Department of Conserva- tion, Columbia. Prins, Rudolph (1972d). Western Kentucky University, Bowling Green. Robison, Henry W. (1982d, 1985b, 1988b, 1989f). Southern Arkansas University, Magnolia. Rodriguez, Gilberto (1989c, 1989d). Instituto Vene- zolano de Investigaciones Cientfficas, Caracas. Shoup, C. S. (1942c, 1947b). Vanderbilt University, Nashville, Tennessee. Villalobos (Figueroa), (1958f, Alejandro 1964a, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1974b, 1981a). Instituto de Biologia, Universidad Nacional Autonoma de México, México. Walton sisters, Lucille (“Miss Lucille’’) and Margaret (“Miss Peggy”) (1957b, 1958d, 1959a, 1959c, 1960a, 1960c, 1961a, 1962b, 1963a, 1963b, 1966c, 1966e, 1967d, 1968b, 1968d, 1970b, 1971le, 1975b, 1976d, 1977f). Danville, Virginia and Mountain Lake Biological Station, Pembroke, Virginia. J. E Fitzpatrick, Jr. (in litt.) notes that ““They were—or at least Hobbs thought of them as—the quintessen- tial “Old Maid Schoolteachers.’ Both were older than he, Miss Lucille by a greater margin .... To the best of my knowledge they met at the Mountain Lake Biological Station where the sisters were sum- mer fixtures. They succumbed to his characteristic charm, and Miss Peggy, “who knew not a thing about crayfishes,”” took great satisfaction in contrib- uting to science by inking the pencil drawings. This seemed to be her contribution to the joint papers.” Whiteman, Mike (1987c, 1991a). Texas Agricultural Extension Service, Lufkin. Word, Benjamin H. (1958e). University of Virginia, Charlottesville. An undergraduate, now a M.D., who did a research project under Hobbs that was never published. At the time of his death Hobbs was pre- paring this report for publication; Hobbs III is pre- paring this final study (see also remarks at end of Hobbs’s bibliography). Zinn, Donald J. (1948c). University of Rhode Island, Kingston. Publications of Horton H. Hobbs, Jr. Here we provide a complete bibliography of Hobbs’s publications, in chronological order, annotated with the names of new taxa in each publication. The citations are cross- referenced to the list of taxa named by Hobbs, given below. If the name of a taxon is given in the title, it is not repeated in the list of taxa named in that article. 1937. Some Florida crawfishes and their habitat dis- tribution. [Abstract]—Proceedings of the Florida Academy of Sciences for 1936 1:154. 1938a. Two new crawfishes from Florida. Cambarus hubbelli, Cambarus acherontis pallidus. [Ab- stract].—Proceedings of the Florida Academy of Sciences 2:90, 91. [Nomina nuda]. 1938b. A new crawfish from Florida.—Journal of the Washington Academy of Sciences 28(2):61—65. Cambarus rogersi. 1940a. Seven new crayfishes of the genus Cambarus from Florida, with notes on other species.—Pro- ceedings of the United States National Museum 89: 387—423. C. hubbelli, C. kilbyi, C. lucifugus ala- chua, C. lucifugus lucifugus, C. pallidus, C. pictus, C. rathbunae. VOLUME 111, NUMBER 3 1940b. A new crayfish from South Carolina.—The Charleston Museum Leaflet 14:3—7. Cambarus lun- Zi. 1941a. A new crayfish from San Luis Potosi, México (Decapoda, Astacidae).—Zoologica, New York 26(1):1—4. Cambarus blandingii cuevachicae. 1941b. Three new Florida crayfishes of the subgenus Cambarus (Decapoda, Astacidae).—The American Midland Naturalist 26(1):110-121. C. byersi, C. cryptodytes, C. floridanus. 1942a. On the first pleopod of the male Cambari (De- capoda, Astacidae).—Proceedings of the Florida Academy of Sciences (for 1940) 5:55-61. 1942b. A generic revision of the crayfishes of the sub- family Cambarinae (Decapoda, Astacidae) with the description of a new genus and species.—The American Midland Naturalist 28(2):334—-357. Trog- locambarus, T. maclanei. 1942c. Hobbs, H. H., Jr., & C. S. Shoup. On the cray- fish collected from the Big South Fork of the Cum- berland River in Tennessee during the summer of 1938.—The American Midland Naturalist 28(3): 634-643. 1942d. The crayfishes of Florida.—University of Flor- ida Publication, Biological Science Series 3(2):179 pp.. pls. 1 (frontispiece), 2—24. Cambarellus schmit- ti, Procambarus apalachicolae, P. bivittatus, P. econfinae, P. escambiensis, P. geodytes, P. latipleu- rum, P. leonensis, P. okaloosae, P. pubischelae, P. Pycnogonopodus, P. pygmaeus, P. rogersi campes- tris, P. r. ochlocknensis, P. seminolae, P. shermani, P. youngi. 1943a. Hobbs, H. H., Jr., & L. J. Marchand. A contri- bution toward a knowledge of the crayfishes of the Reelfoot Lake area.—Journal of the Tennessee Academy of Science 18(1):6—35. 1943b. Two new crayfishes from the panhandle of Florida (Decapoda, Astacidae).—Proceedings of the Florida Academy of Sciences 6(1):49-—58. Note by Hobbs given in footnote (p. 56): ““Due to inadver- tent delay in publication of this volume diagnoses of these species [Procambarus leonensis, P. pyc- nogonopodus| appeared earlier in “The Crayfishes of Florida’ (Hobbs: Univ. Fla. Pub. Biol. Series 3(2): 114-115, 117) and thus actually constitute the orig- inal descriptions.” 1943c. Two new crayfishes of the genus Procambarus from Mexico (Decapoda, Astacidae).—Lloydia 6: 198-206. P. rodriguezi, P. toltecae. 1944. Notes on the subterranean waters of the Florida Peninsula with particular reference to their crusta- cean fauna.—The Biologist 26(1&2):6-8. 1945a. Notes on the first pleopod of the male Cam- barinae (Decapoda, Astacidae).—Quarterly Journal of the Florida Academy of Sciences 8(1):67—70. 1945b. The subspecies and intergrades of the Florida burrowing crayfish, Procambarus rogersi (Hobbs).—Journal of the Washington Academy of 487 Sciences 35(8):247—260. Note by Hobbs given in footnote (p. 260): “This paper was originally ac- cepted for publication in the Proceedings of the United States National Museum, and it was cited as ‘In press’ in my Crayfishes of Florida (Hobbs, 1942). Wartime restrictions, however, so delayed publication by the Museum that the manuscript was withdrawn and submitted to this JOURNAL in order that the full descriptions of the two new subspecies of Procambarus rogersi might appear more prompt- ly.” 1945c. Two new species of crayfishes of the genus Cambarellus from the Gulf coastal states, with a key to the species of the genus (Decapoda, Astacidae).— The American Midland Naturalist 34(2):466—474. C. diminutus, C. puer. 1947a. Two new crayfishes of the genus Procambarus from Georgia, with notes on Procambarus pubes- cens (Faxon) (Decapoda, Astacidae).—Quarterly Journal of the Florida Academy of Sciences 9(1):1— 18. P. enoplosternum, P. litosternum. 1947b. Hobbs, H. H., Jr, & C. S. Shoup. Two new crayfishes (Decapoda, Astacidae) from the Obey River drainage in Tennessee.—Journal of the Ten- nessee Academy of Science 22(2):138—-145. Cam- barus obeyensis, C. parvoculus. 1947c. A key to the crayfishes of the Pictus subgroup of the genus Procambarus, with the description of a new species from South Carolina.—The Florida Entomologist 30(3):25—31. P. lepidodactylus. 1947d. A preliminary report on the crayfishes of Vir- ginia. [Abstract].—Proceedings of the Virginia Academy of Science 1946—1947:72. 1948a. On the crayfishes of the Limosus section of the genus Orconectes (Decapoda: Astacidae).—Journal of the Washington Academy of Sciences 38(1):14— 21. O. shoupi. 1948b. Two new crayfishes of the genus Orconectes from Arkansas, with a key to the species of the Hy- las group (Decapoda: Astacidae).—The American Midland Naturalist 39(1):139-150. O. leptogono- podus, O. marchandi. 1948c. Hobbs, H. H., Jr, & D. J. Zinn. Crayfish in southern Nevada.—Science 107(2780):369. 1948d. The crayfish genus Cambarellus in the United States. [Abstract].—Proceedings of the Virginia Academy of Science 1948:88. 1948e. A new crayfish of the genus Orconectes from southern Tennessee (Decapoda, Astacidae).—Pro- ceedings of the Biological Society of Washington 61:85—91. O. wrighti. 1948f. A new crayfish of the genus Cambarus from Texas, with notes on the distribution of Cambarus fodiens (Cottle).—Proceedings of the United States National Museum 98:223-—231. C. hedgpethi. 1949a. The cave crayfishes of North America. [Ab- stract].—Journal of the Tennessee Academy of Sci- ence 24(3):170. 488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1949b. Hobbs, H. H., Jr, & C. E. Parish. Notes on the life history of a Virginia crayfish. [Abstract].—Pro- ceedings of the Virginia Academy of Science 1948— 1949:97. 1949c. Observations on the emergence of a stonefly of the genus Taeniopteryx in Virginia. [Abstract].— Proceedings of the Virginia Academy of Science 1948—1949:101. 1949d. A new crayfish of the genus Orconectes from the Nashville Basin in Tennessee, with notes on the range of Orconectes compressus (Faxon) (Decapo- da, Astacidae).—Proceedings of the Biological So- ciety of Washington 62:17—25. O. rhoadesi. 1950a. A new crayfish of the genus Cambarellus from Texas (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 63:89—94. C. ni- nae. 1950b. A new crayfish of the genus Procambarus from Oklahoma and Arkansas (Decapoda, Astacidae).— Journal of the Washington Academy of Sciences 40(6):194—198. P. tenuis. 1950c. Observations on the ecological distribution of three Virginia crayfishes. [Abstract]—The Virginia Journal of Science for 1949-1950, new series 1(4): 349. 1951a. A new crayfish of the genus Orconectes from southeastern Virginia (Decapoda, Astacidae).—The Virginia Journal of Science, new series 2(2):122— 128. O. virginiensis. 1951b. A new crayfish of the genus Procambarus from Louisiana, with a key to the species of the Spiculifer group.—Journal of the Washington Academy of Sci- ences 41(8):272—276. P. penni. 1952a. A new crayfish from Alabama, with notes on Procambarus lecontei (Hagen).—Proceedings of the United States National Museum 102:209-219. P. verrucosus. 1952b. Hobbs, H. H., Jr., & W. H. Massmann. The river shrimp, Macrobrachium ohione (Smith), in Virginia—The Virginia Journal of Science, new se- ries 3(3):206, 207. 1952c. A new crayfish of the genus Procambarus from Georgia with a key to the species of the Clarkii sub- group.—Quarterly Journal of the Florida Academy of Sciences 15(3):165—-174. P. howellae. 1952d. A new albinistic crayfish of the genus Cam- barus from southern Missouri with a key to the al- binistic species of the genus (Decapoda, Astaci- dae)—The American Midland Naturalist 48(3): 689-693. C. hubrichti. 1952e. A preliminary report on the crayfishes of the Atlantic Slope from New Brunswick to South Car- olina. [Abstract]——The Virginia Journal of Science, new series 3(4):295. 1953a. Two new crayfishes from the Highland Rim in Tennessee (Decapoda, Astacidae).—Journal of the Tennessee Academy of Science 28(1):20—27 [also published in Report of the Reelfoot Lake Biological Station, vol. 17, 1953]. Cambarus brachydactylus, C. friaufi. 1953b. Hobbs, H. H., Jr, & C. H. Page. Additional records of the occurrence of the freshwater jellyfish, Craspedacusta sowerbii, in Virginia.—The Virginia Journal of Science, new series 4(3):137. 1953c. The epizootic associates of the crayfishes of the New River system with particular reference to the ostracods. [Abstract].—Journal of the Tennessee Academy of Science 28(3):180, 181. 1953d. A new crayfish of the genus Procambarus from Alabama and Florida (Decapoda, Astacidae).—Pro- ceedings of the Biological Society of Washington 66:173-178. P. suttkusi. 1953e. On the ranges of certain crayfishes of the Spi- culifer group of the genus Procambarus, with the description of a new species (Decapoda: Astaci- dae).—Journal of the Washington Academy of Sci- ences 43(12):412—417. P. raneyi. 1954a. Apparent competition between two groups of crayfishes in the southeastern states. [Abstract].— The Virginia Journal of Science, new series 4(4): 230. 1954b. A new crayfish from the upper coastal plain of Georgia (Decapoda, Astacidae).—Quarterly Journal of the Florida Academy of Sciences 17(2):110—118. Procambarus truculentus. 1954c. A redescription of Procambarus ruthveni (Pearse) from La Laja Creek at Cuatatotolapam, Ve- racruz, Mexico (Decapoda, Astacidae).—Occasional Papers of the Museum of Zoology, University of Michigan 559:1-5. 1954d. Studies on the geographic distribution of the crayfishes of the genus Procambarus. [Abstract].— Journal of the Tennessee Academy of Science 29(3): 181. 1954e. Notes on the evolution of the Longulus group of the crayfish genus Cambarus. [Abstract]—The Virginia Journal of Science, new series 5(4):261. 1955a. A new crayfish of the genus Cambarus from Mississippi.—Proceedings of the Biological Society of Washington 65:95—100. C. cristatus. 1955b. Ostracods of the genus Entocythere from the New River system in North Carolina, Virginia, and West Virginia.—Transactions of the American Mi- croscopical Society 74(4):325—333. E. daphnioides, E. runki. 1955c. A tendency towards cyclic dimorphism in fe- male crayfishes. [Abstract]—The Virginia Journal of Science, new series 6(4):248. 1955d. Two crayfish highways to Florida. [Ab- stract]_—Association of Southeastern Biologists Bulletin 2(1):7. 1956a. A new crayfish of the genus Cambarus from North Carolina and South Carolina (Decapoda, As- tacidae)—Journal of the Elisha Mitchell Scientific Society 72(1):61—67. C. reduncus. 1956b. A new crayfish of the Extraneus section of the VOLUME 111, NUMBER 3 genus Cambarus with a key to the species of the section (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 69:115-121. C. spicatus. 1956c. A new crayfish of the genus Procambarus from South Carolina (Decapoda: Astacidae).—Journal of the Washington Academy of Sciences 46(4):117— 121. P. echinatus. 1956d. Hobbs, H. H., Jr., & H. W. Freeman. The deca- pod crustaceans of the Wateree River system in North Carolina and South Carolina. [Abstract].— Association of Southeastern Biologists Bulletin 3(1):10. 1956e. Hart, C. W., & H. H. Hobbs, Jr. The crayfish of the Lower Flint-Chattahoochee River system. [Abstract]—The Virginia Journal of Science, new series 7(4):292. 1957a. Observaciones acerca de las especies del gé- nero Entocythere (Crustaceos, Ostracodos) de Cuba.—Anales del Instituto de Biologia, Universi- dad Nacional Aut6noma de México 27(2):431—436. E. hamata. 1957b. Hobbs, H. H., Jr, & M. Walton. Three new crayfishes from Alabama and Mississippi (Decapo- da: Astacidae).—Tulane. Studies in Zoology 5(3): 39-52. Procambarus hybus, P. jaculus, P. mancus. 1958a. The evolutionary history of the Pictus group of the crayfish genus Procambarus (Decapoda, Asta- cidae).—Quarterly Journal of the Florida Academy of Sciences 2(1):71—91. 1958b. Two new crayfishes of the genus Procambarus from South Carolina.—Journal of the Washington Academy of Sciences 48(5):160—168. P. ancylus, P. hirsutus. 1958c. Two new crayfishes of the genus Procambarus from South Carolina and Georgia.—Notulae Natu- rae, Academy of Natural Sciences of Philadelphia 307:1—10, pls. 1, 2. P. chacei, P. epicyrtus. 1958d. Hobbs, H. H., Jr, & M. Walton. Procambarus pearsei plumimanus, a new crayfish from North Car- olina (Decapoda, Astacidae).—Journal of The Eli- sha Mitchell Scientific Society 74(1):7—12. 1958e. Word, B. H., & H. H. Hobbs, Jr. Observations on the testis of the crayfish Cambarus montanus ac- uminatus Faxon.—Transactions of the American Microscopical Society 77(4):435—450. 1958f. Hobbs, H. H., Jr, & A. Villalobos. The exo- skeleton of a freshwater crab as a microhabitat for several invertebrates. [Abstract].—The Virginia Journal of Science, new series 9(4):395, 396. 1958g. General Zoology. Tracey I. Storer & Robert L. Usinger. [Book review].—American Institute of Bi- ological Sciences Bulletin 8:43. 1958h. Penn, G. H., & H. H. Hobbs, Jr. A contribution toward a knowledge of the crawfishes of Texas (De- capoda, Astacidae)—The Texas Journal of Science 10(4):452-—483. 1959a. Hobbs, H. H., Jr., & M. Walton. A new crayfish 489 of the genus Procambarus from Alabama (Decap- oda, Astacidae).—Proceedings of the Biological So- ciety of Washington 72:39—44. P. lewisi. 1959b. Hobbs, H. H., Jr., & C. W. Hart, Jr. The fresh- water decapod crustaceans of the Apalachicola drainage system in Florida, Southern Alabama, and Georgia.—Bulletin of the Florida State Museum, Bi- ological Series 4(5):145—191. Procambarus rogersi expletus. 1959c. Walton, M., & H. H. Hobbs, Jr. Two new eye- less ostracods of the genus Entocythere from Flori- da.—The Quarterly Journal of the Florida Academy of Sciences 22(2):114—120. E. ambophora, E. luci- fuga. 1959d. Hobbs, H. H., Jr., E A. Chace, Jr., J. G. Mackin, L. Hubricht, & A. H. Banner. Malacostraca. Pp. 889-901 in H. B. Ward & G. C. Whipple (W. T. Edmondson, editor), Freshwater Biology, 2nd edi- tion, Wiley, New York, 1248 pp. [crayfishes, pp. 882-898]. 1960a. Hobbs, H. H., Jr, & M. Walton. Three new ostracods of the genus Entocythere from the Hiwas- see drainage system in Georgia and Tennessee.— Journal of the Tennessee Academy of Science 35(1): 17-23. E. cyma, E. mecoscapha, E. simondsi. 1960b. Hobbs, H. H., Jr, & T. C. Barr, Jr. The genus Cambarus. The origins and affinities of the troglob- itic crayfishes of North America (Decapoda, Asta- cidae), I—The American Midland Naturalist 64(1): 12—33. C. jonesi. 1960c. Hobbs, H. H., Jr., & M. Walton. A new crayfish of the genus Procambarus from southern Alabama (Decapoda, Astacidae).—Proceedings of the Biolog- ical Society of Washington 73:123—129. P. lophotus. 1961a. Hobbs, H. H., Jr., & M. Walton. Additional new ostracods from the Hiwassee drainage system in Georgia, North Carolina, and Tennessee.—Transac- tions of The American Microscopical Society 80(4): 379-384. Entocythere falcata, E. hiwasseensis. 1961b. Hart, C. W., Jr, & H. H. Hobbs, Jr. Eight new troglobitic ostracods of the genus Entocythere (Crustacea, Ostracoda) from the eastern United States.—Proceedings of The Academy of Natural Sciences of Philadelphia 113(8):173-185. E. arcu- ata, E. barri, E. pholetera, E. prionata, E. steevesi, E. tuberosa, E. ungulata, E. xania. 1962a. Hobbs, H. H., Jr, & H. H. Hobbs Ill. A new crayfish of the genus Cambarus from Georgia (De- capoda, Astacidae).—Proceedings of the Biological Society of Washington 75:41—45. C. conasaugaen- SiS. 1962b. Hobbs, H. H., Jr, & M. Walton. New ostracods of the genus Entocythere from the Mountain Lake Region, Virginia (Ostracoda, Entocytheridae).—The Virginia Journal of Science, new series 13(2):42—48. E. asceta, E. chalaza, E. phyma. 1962c. Notes on the affinities of the members of the Blandingii section of the crayfish genus Procam- 490 barus (Decapoda, Astacidae).—Tulane Studies in Zoology 9(5):273—293. 1962d. Hobbs, H. H., Jr., & J. FE Fitzpatrick, Jr. A new crayfish of the Propinquus group of the genus Or- conectes from the Ohio system drainage in West Virginia (Decapoda: Astacidae).—Proceedings of the Biological Society of Washington 75:207—214. O. propinquus erismophorous. 1962e. La presencia de Procambarus clarkii (Girard) en los estados de Chihuahua y Sonora, México (De- capoda, Astacidae).—Anales del Instituto de Biol- ogia, Universidad Nacional Aut6noma de México 33(1 &2):273—276. 1963a. Hobbs, H. H., Jr., & M. Walton. Three new ostracods (Ostracoda, Entocytheridae) from the Duck River drainage in Tennessee.—The American Midland Naturalist 69(2):456—461. Ankylocythere hyba, Dactylocythere xystroides, Uncinocythere zancla. 1963b. Hobbs, H. H., Jr., & M. Walton. Four new spe- cies of the genus Donnaldsoncythere (Ostracoda, Entocytheridae) from Virginia with a key to the spe- cies of the genus.—Transactions of the American Microscopical Society 82(4):363—-370. D. ardis, D. ileata, D. scalis, D. truncata. 1963c. Florida crayfishes.—The Quipu 1(2):8. 1964a. Hobbs, H. H., Jr, & A. Villalobos. Los cam- barinos de Cuba.—Anales del Instituto de Biologia, Universidad Nacional Autonoma de México 84(1 &2):307—366. Procambarus niveus. 1964b. Hobbs, H. H., Jr, & M. S. Bedinger. A new troglobitic crayfish of the genus Cambarus (Decap- oda, Astacidae) from Arkansas with a note on the range of Cambarus cryptodytes Hobbs.—Proceed- ings of the Biological Society of Washington 77:9— 15. C. zophonastes. 1964c. A new cave-dwelling crayfish from the Green- brier drainage system, West Virginia (Decapoda, As- tacidae).—Proceedings of the Biological Society of Washington 77:189—194. Cambarus nerterius. 1965a. Bedinger, M. S., & H. H. Hobbs, Jr. Observa- tions of a new troglobitic crayfish (with notes on the distribution of troglobitic crayfishes in the Ozark Region).—National Speleological Society, Bulletin 27(3):93-96. 1965b. A new crayfish of the genus Cambarus from Tennessee with an emended definition of the genus (Decapoda, Astacidae).—Proceedings of the Biolog- ical Society of Washington 78:265—-273. C. pristin- us. 1966a. Astacus oreganus Randall, 1840 (Crustacea, Decapoda): proposed suppression under the Plenary Powers. Z.N.(S.) 1727.—Bulletin of Zoological No- menclature 22(5&6):351—354. 1966b. A new crayfish from Alabama with observa- tions on the Cristatus section of the genus Cambarus (Decapoda, Astacidae).—Proceedings of the Biolog- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ical Society of Washington 79:109-116. C. promi- nens. 1966c. Hobbs, H. H., Jr, & M. Walton. Orconectes juvenilis (Hagen) in Mountain Lake, Virginia: an unplanned experiment in interspecific competition (Decapoda, Astacidae)—The Virginia Journal of Science, new series 17(3):136—140. 1966d. Hobbs, H. H., Jr, & C. W. Hart, Jr. On the entocytherid ostracod genera Ascetocythere, Plec- tocythere, Phymocythere (gen. nov.), and Cymocy- there, with description of new species.—Proceed- ings of The Academy of Natural Sciences of Phil- adelphia 118(2):35-61. A. coryphodes, A. cosmeta, A. didactylata, A. hoffmani, A. hyperoche, A. my- xoides, A. ozalea, A. sclera, C. gonia, Plectocythere Johnsonae. 1966e. Hobbs, H. H., Jr., & M. Walton. A new genus and six new species of entocytherid ostracods (Os- tracoda, Entocytheridae).—Proceedings of the Unit- ed States National Museum 119:1—12. Dactylocy- there brachystrix, D. pachysphyrata, D. cayugaen- sis, Entocythere kanawhaensis, Thermastrocythere, T. harti, Unicinocythere stubbsi. 1966f. An illustrated key to the species of the genus Ankylocythere with a description of a new species from Louisiana (Ostracoda, Entocytheridae).—The Proceedings of the Louisiana Academy of Sciences 29:67-75. A. harmani. 1967a. A new genus and three new species of ostra- cods with a key to genus Dactylocythere (Ostracoda: Entocytheridae).—Proceedings of the United States National Museum 122:1—10. D. jeanae, D. phoxa, Ornithocythere, O. waltonae. 1967b. The current status of the crayfishes listed by Girard (1852) in his “‘A revision of the North Amer- ican Astaci ...’’ (Decapoda, Astacidae).—Crusta- ceana 12:124—132. 1967c. Hobbs, H. H., Jr, & E O. Perkins. A new bur- rowing crayfish from North Carolina (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 80:141—146. Cambarus catagius. 1967d. Hobbs, H. H. Jr, P. C. Holt, & M. Walton. The crayfishes and their epizootic ostracod and bran- chiobdellid associates of the Mountain Lake, Vir- ginia, Region.—Proceedings of the United States National Museum 123:1—84. 1967e. A new crayfish from Alabama caves with notes on the origin of the genera Orconectes and Cam- barus (Decapoda: Astacidae).—Proceedings of the United States National Museum 123:1—17. Procam- barus pecki. 1968a. Two new crayfishes of the genus Cambarus from Georgia, Kentucky, and Tennessee (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 81:261—274. C. halli, C. sphenoides. 1968b. Hobbs, H. H., Jr., BR C. Holt, & M. Walton. The crayfishes and their epizootic ostracod and bran- chiobdellid associates of the Mountain Lake, Vir- VOLUME 111, NUMBER 3 ginia, region. [Abstract]—The Virginia Journal of Science, new series 19(3):178. 1968c. Crustacea: Malacostraca. Pp. K1—-K36 in E K. Parrish, Keys to water quality indicative organisms (southeastern United States). Federal Water Pollu- tion Control Administration, U.S. Department of In- terior. 1968d. Hobbs, H. H., Jr., & M. Walton. New entocy- therid ostracods from the southern United States.— Proceedings of the Academy of Natural Sciences of Philadelphia 120(6):237—252. Ascetocythere batchi, Dactylocythere cooperorum, D. prinsi, Entocythere reddelli, Harpagocythere tertius, Litocythere, L. lu- cileae, Uncinocythere warreni. 1969a. E A. Chace, Jr., & H. H. Hobbs, Jr. The fresh- water and terrestrial decapod crustaceans of the West Indies with special reference to Dominica.— United States National Museum Bulletin 292:258 Pp. 1969b. Procambarus villalobosi, un nuevo cambarino de San Luis Potosi, México (Decapoda, Astaci- dae).—Anales del Instituto de Biologia, Universidad Nacional Aut6énoma de México, Serie Ciencias del Mar y Limnologia, 38(1)(for 1967):41—46. 1969c. On the distribution and phylogeny of the cray- fish genus Cambarus. Pp. 93-178 in P. C. Holt, R. L. Hoffman, & C. W. Hart, Jr, eds. The distribu- tional history of the biota of the southern Appala- chians, Part I: Invertebrates. Virginia Polytechnic In- stitute, Research Division Monograph 1:295 pp. Av- iticambarus, Barbicambarus, Depressicambarus, Erebicambarus, Fallicambarus, Jugicambarus, Lacunicambarus, Veticambarus. 1969d. Hobbs, H. H., Jr, & E. T. Hall, Jr. New cray- fishes from Georgia (Decapoda Astacidae).—Pro- ceedings of the Biological Society of Washington 82:281—294. Cambarus howardi, C. unestami. 1969e. Two new species of the crayfish genus Procam- barus (Decapoda, Astacidae) with keys to the mem- bers of the Spiculifer group.—Proceedings of the Biological Society of Washington 82:329-348. P. elegans, P. gibbus. 1970a. Hobbs, H. H., Jr., & J. E Fitzpatrick, Jr. A new crayfish of the genus Fallicambarus from Tennessee (Decapoda, Astacidae).—Proceedings of the Biolog- ical Society of Washington 82:829—836. F. hortoni. 1970b. Hobbs, H. H., Jr., & M. Walton. New entocy- therid ostracods from Tennessee and Virginia.—Pro- ceedings of the Biological Society of Washington 82:851—863. Ascetocythere holti, Dactylocythere en- oploholea, D. myura, D. spinata. 1970c. Hobbs, H. H., Jr, & H. H. Hobbs III. New entocytherid ostracods with a key to the genera of the subfamily Entocytherinae.—Smithsonian Con- tributions to Zoology 47:19 pp. Ascetocythere lita, Dactylocythere coloholca, D. macroholca, D. pug- Hiaticambarus, Puncticambarus, 491 hae, Entocythere tyttha, Geocythere nessoides, Lor- docythere, L. petersi. 1970d. A new crayfish from the Nashville Basin, Ten- nessee.—Proceedings of the Biological Society of Washington 83:161—169. Cambarus gentryi. 1970e. New crayfishes of the genus Cambarus from Tennessee and Georgia (Decapoda, Astacidae).— Proceedings of the Biological Society of Washing- ton 83:241—259. C. bouchardi, C. cymatilis. 1971a. A new crayfish of the genus Procambarus from Mississippi (Decapoda: Astacidae).—Proceedings of the Biological Society of Washington 83:459—468. P. fitzpatricki. 1971b. The entocytherid ostracods of Mexico and Cuba.—Smithsonian Contributions to Zoology 81: 55 pp. Ankylocythere maya, A. toltecae, A. villalo- bosi, Uncinocythere zaruri. 1971c. New crayfishes of the genus Procambarus from Alabama and Texas (Decapoda, Astacidae).—Pro- ceedings of the Biological Society of Washington 84:81—94. P. capillatus, P. texanus. 1971d. Fitzpatrick, J. E, Jr., & H. H. Hobbs, Jr. A new crawfish of the Spiculifer group of the genus Pro- cambarus (Decapoda, Astacidae) from central Mis- Sissipp1.—Proceedings of the Biological Society of Washington 84:95—-102. P. lylei. 1971le. Walton, M., & H. H. Hobbs, Jr. The distribution of certain entocytherid ostracods on their crayfish hosts.—Proceedings of the Academy of Natural Sci- ences of Philadelphia 123(4):87—103. 1971f. A new troglobitic crayfish from Florida.— Quarterly Journal of the Florida Academy of Sci- ences 34(2):114—-124. Procambarus milleri. 1972a. The subgenera of the crayfish genus Procam- barus (Decapoda: Astacidae).—Smithsonian Contri- butions to Zoology 117:22 pp. Acucauda, Austro- cambarus, Capillicambarus, Hagenides, Leconti- cambarus, Lonnbergius, Mexicambarus, Pennides, Remoticambarus, Scapulicambarus, Tenuicamba- rus, Villalobosus. 1972b. Hobbs, H. H., Jr., & T. C. Barr, Jr. Genus Or- conectes. Origins and affinities of the troglobitic crayfishes of North America (Decapoda, Astacidae), II.—Smithsonian Contributions to Zoology 105:84 pp. O. incomptus. 1972c. Hobbs, H. H., Jr, & D. B. Means. Two new troglobitic crayfishes (Decapoda, Astacidae) from Florida.—Proceedings of the Biological Society of Washington 84:393—409. Procambarus horsti, P. orcinus. 1972d. Prins, R., & H. H. Hobbs, Jr. A new crayfish of the subgenus Puncticambarus from the Savannah River drainage with notes on Cambarus (P.) rebur- rus Prins (Decapoda, Astacidae).—Proceedings of the Biological Society of Washington 84:411—420. Cambarus chaugaensis. 1972e. Hobbs, H. H., Jr, & M. R. Cooper. A new troglobitic crayfish from Oklahoma (Decapoda: As- 492 tacidae).—Proceedings of the Biological Society of Washington 85:49—56. Cambarus tartarus. 1972f. Hobbs, H. H., Jr, & E. T. Hall, Jr. A new cray- fish from the Tallapoosa River in Georgia (Decap- oda: Astacidae).—Proceedings of the Biological So- ciety of Washington 85:151—161. Cambarus engli- shi. 1972g. Crayfishes (Astacidae) of North and Middle America.—Biota of Freshwater Ecosystems, Identi- fication Manual No. 9:173 pp. United States Envi- ronmental Protection Agency, Water Pollution Con- trol Research Series. 1973a. Hobbs, H. H., Jr, & R. W. Bouchard. A new crayfish from the Cumberland River system with notes on Cambarus carolinus (Erichson).—Proceed- ings of the Biological Society of Washington 86:41— 68. Cambarus cumberlandensis. 1973b. Three new troglobitic decapod crustaceans from Oaxaca, Mexico. In R. E. Mitchell & J. Red- dell, eds., Studies on the cavernicole fauna of Mex- ico and adjacent regions.—Bulletin, Association for Mexican Cave Studies 5:25—38. Neopalaemon, N. nahuatlus, Procambarus oaxacae oaxacae, P. oax- acae reddelli. 1973c. Hobbs, H. H., Jr, & H. H. Hobbs III. The genus Sphaeromicola (Ostracoda, Entocytheridae) in Mex- ico. In R. E. Mitchell & J. Reddell, eds. Studies on the cavernicole fauna of Mexico and adjacent regions.—Bulletin, Association for Mexican Cave Studies 5:39—42. S. coahuiltecae. 1973d. Two new troglobitic shrimp (Decapoda: Al- pheidae and Palaemonidae) from Oaxaca, Mexico. In R. E. Mitchell & J. Reddell, eds., Studies on the cavernicole fauna of Mexico and adjacent regions.—Bulletin, Association for Mexican Cave Studies 5:73-80. Alpheopsis stygicola, Macrobra- chium villalobosi. 1973e. New species and relationships of the members of the genus Fallicambarus.—Proceedings of the Biological Society of Washington 86:461—482. Creaserinus, F. jeanae, F. spectrum. 1974a. Synopsis of the families and genera of cray- fishes (Crustacea: Decapoda).—Smithsonian Contri- butions to Zoology 164:32 pp. 1974b. Villalobos Figueroa, A., & H. H. Hobbs, Jr. Three new crustaceans from La Media Luna, San Luis Potosi, Mexico.—Smithsonian Contributions to Zoology 174:18 pp. Ankylocythere barbouri, Palae- monetes lindsayi, Procambarus roberti. 1974c. A checklist of the North and Middle American crayfishes (Decapoda: Astacidae and Cambari- dae).—Smithsonian Contributions to Zoology 166: 161 pp. 1974d. Hobbs, H. H., Jr, & E. T. Hall, Jr. Crayfishes (Decapoda: Astacidae). Pp. 195-214 in C. W. Hart, Jr. & S. L. H. Fuller, eds., Pollution ecology of fresh- water invertebrates. 389 pp. Academic Press, New York. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1975a. Key to the troglobitic crayfishes of Florida. Pp. 14, 15 in K. Relyea & B. Sutton, A new troglobitic crayfish of the genus Procambarus from Florida (Decapoda: Astacidae).—Tulane Studies in Zoology and Botany 19(1&2):8—-16. 1975b. Hobbs, H. H., Jr., & M. Walton. New entocy- therid ostracods from Tennessee with a key to the species of the genus Ascetocythere.—Proceedings of the Biological Society of Washington 88:5—20. A. bouchardi, A. pseudolita, A. triangulata, A. veruta, Dactylocythere crena, D. scissura, Psittocythere, P. psitta. 1975c. New crayfishes (Decapoda: Cambaridae) from the southern United States and Mexico.—Smithson- ian Contributions to Zoology 201:34 pp. Fallicam- barus caesius, F. danielae, Procambarus clemmeri, P. geminus, P. marthae, P. medialis, P. xochitlanae. 1976a. Adaptations and convergence in North Ameri- can crayfishes. Pp. 541-551 in J. W. Avault, Jr., ed., Freshwater crayfish.—Papers from the Second In- ternational Crayfish Symposium, Baton Rouge, Lou- isiana, USA, 1974. 676 pp. Louisiana State Univer- sity. 1976b. Bouchard, R. W., & H. H. Hobbs, Jr A new subgenus and two new species of crayfishes of the genus Cambarus (Decapoda: Cambaridae) from the southern United States—Smithsonian Contributions to Zoology 224:15 pp. C. cracens, C. nodosus, Ex- ilicambarus. 1976c. Hobbs, H. H., Jr., & D. S. Lee. A new troglob- itic crayfish (Decapoda, Cambaridae) from penin- sular Florida.—Proceedings of the Biological Soci- ety of Washington 89:383-—391. Procambarus franzi. 1976d. Hobbs, H. H., Jr, & M. Walton. New entocy- therid ostracods from Kentucky and Tennessee.— Proceedings of the Biological Society of Washing- ton 89:393—404. Ascetocythere riopeli, Dactylocy- there apheles, D. brachydactylus, D. demissa. 1976e. Hobbs Ill, H. H., & H. H. Hobbs, Jr. On the troglobitic shrimps of the Yucatan Peninsula, Mex- ico (Decapoda: Atyidae and Palaemonidae).— Smithsonian Contributions to Zoology 240:23 pp. Typhlatya campecheae, T. mitchelli. 1977a. The crayfish Bouchardina robisoni, a new ge- nus and species (Decapoda, Cambaridae) from southwestern Arkansas.—Proceedings of the Bio- logical Society of Washington 89:733-742. 1977b. Hobbs, H. H., Jr., & D. J. Peters. The entocy- therid ostracods of North Carolina—Smithsonian Contributions to Zoology 247:73 pp. Aphelocythere, A. acuta, Dactylocythere isabelae, D. peedeensis, Donnaldsoncythere leptodrilus, Entocythere costata, Harpagocythere baileyi. 1977c. Hobbs, Jr., H. H., H. H. Hobbs III, & M. A. Daniel. A review of the troglobitic decapod crusta- ceans of the Americas.—Smithsonian Contributions to Zoology 244:183 pp. 1977d. Manning, R. B., & H. H. Hobbs, Jr. Decapoda. VOLUME 111, NUMBER 3 Pp 157-162 in S. H. Hurlbert, ed., Biota Acuatica de Sudameérica Austral. San Diego State University, San Diego, California, 342 pp. 1977e. A new crayfish (Decapoda: Cambaridae) from San Luis Potosi, Mexico.—Proceedings of the Bio- logical Society of Washington 90(2):412—419. Pro- cambarus strenthi. 1977f. Hobbs, H. H., Jr., & M. Walton. New entocy- therid ostracods of the genus Dactylocythere.—Pro- ceedings of the Biological Society of Washington 90(3):600—614. D. astraphes, D. corvus, D. promi- nula, D. spinescens. 1978. Cave-inhabiting crayfishes of Chiapas, Mexico (Decapoda: Cambaridae). Jn Subterranean Fauna of Mexico, part III, Further results of the Italian zoo- logical missions to Mexico sponsored by the Na- tional Academy of Lincei (1973 and 1975).—Ac- cademia Nazionale dei Lincei, Problemi Attuali di Scienza e di Cultura, Sezione: Missioni ed Esplor- azioni—I [for 1977] 171:197—206. Procambarus sbordonii. 1979a. Hobbs, H. H., Jr., & D. J. Peters. A substitute name for the homonym Aphelocythere Hobbs & Pe- ters (Ostracoda: Entocytheridae)—Proceedings of the Biological Society of Washington 91(4):1037. Waltoncythere. 1979b. A new crayfish from the Ouachita River Basin in Arkansas (Decapoda: Cambaridae).—Proceedings of the Biological Society of Washington 92(4):804— 811. Procambarus reimeri. 1980a. Atya gabonensis (Decapoda Atyidae) in the Western Hemisphere.—Crustaceana 38(1):111, 112. 1980b. New dwarf crayfishes (Decapoda: Cambaridae) from Mexico and Florida.—Proceedings of the Bi- ological Society of Washington 93(1):194-207. Cambarellus blacki, C. chihuahuae. 1980c. Cooper, M. R., & H. H. Hobbs, Jr. New ‘and little-known crayfishes of the virilis section of genus Orconectes (Decapoda: Cambaridae) from the southeastern United States.—Smithsonian Contri- butions to Zoology 320:44 pp. O. chickasawae, O. cooperi, O. holti. 1980d. A new pseudothelphusid crab from the State of Jalisco, Mexico.—Proceedings of the Biological So- ciety of Washington 93(2):357—361. Pseudothelphu- Sa Seiferti. 1981la. Villalobos Figueroa, A., & H. H. Hobbs, Jr. A new dwarf crayfish from the Pacific versant of Mex- ico (Decapoda: Cambaridae).—Proceedings of the Biological Society of Washington 94(2):492—502. Cambarellus prolixus. 1981b. The crayfishes of Georgia.—Smithsonian Con- tributions to Zoology 318:549 pp. Cambarus acan- thura, C. coosae, C. coosawattae, C. fasciatus, C. georgiae, C. harti, C. hiwasseensis, C. manningi, C. parrishi, C. reflexus, C. scotti, C. speciosus, C. stri- gosus, C. truncatus, Distocambarus, Procambarus 493 caritus, P. devexus, P. petersi, P. pubischelae defi- ciens, P. talpoides. 1982a. A new crayfish (Decapoda: Cambaridae) from the State of Puebla, Mexico, with new locality re- cords for Procambarus (Villalobosus) xochitlanae and entocytherid ostracod symbionts.—Association for Mexican Cave Studies Bulletin 8:39—44 (also Texas Memorial Museum Bulletin 28:39—44). Pro- cambarus cuetzalanae. 1982b. Hobbs, H. H., Jr, & A. G. Grubbs. Description of a new troglobitic crayfish from Mexico and a list of Mexican crayfishes reported since the publication of the Villalobos Monograph (1955) (Decapoda, Cambaridae).—Association for Mexican Cave Stud- ies Bulletin 8:45—SO (also Texas Memorial Museum Bulletin 28:45—-50). Procambarus xilitlae. 1982c. Hobbs, H. H., Jr., & D. J. Peters. The entocy- therid ostracod fauna of northern Georgia.—Pro- ceedings of the Biological Society of Washington 95(2):297-3 18. 1982d. Hobbs, H. H., Jr, & H. W. Robison. A new crayfish of the genus Procambarus from southwest- ern Arkansas.—Proceedings of the Biological So- ciety of Washington 95(3):545—-553. P. parasimu- lans. 1982e. Hobbs, H. H., Jr., & C. W. Hart, Jr. The shrimp genus Atya (Decapoda: Atyidae).—Smithsonian Contributions to Zoology 364:143 pp. A. brachyr- hinus. 1982f. On the distribution of the genus Procambarus. Pp. 2, 3 [Abstract] in J. E Payne, ed., Crayfish dis- tribution patterns, American Society of Zoologists. 1983a. Franz, R., & H. H. Hobbs, Jr. Procambarus (Ortmannicus) leitheuseri, new species, another troglobitic crayfish (Decapoda: Cambaridae) from peninsular Florida.—Proceedings of the Biological Society of Washington 96(2):323—332. 1983b. The African shrimp genus Potamalpheops in Mexico (Decapoda: Alpheidae).—Crustaceana 44(2):221—224. 1983c. Hobbs, H. H., Jr., & P H. Carlson. Distocam- barus (Decapoda: Cambaridae) elevated to generic rank, with an account of D. crockeri, new species, from South Carolina.—Proceedings of the Biologi- cal Society of Washington 96(3):420—428. 1983d. Distocambarus (Fitzcambarus) carlsoni, a new subgenus and species of crayfish (Decapoda: Cam- baridae) from South Carolina.—Proceedings of the Biological Society of Washington 96(3):429—439. 1983e. Hobbs, H. H., Jr., & A. C. McClure. On a small collection of entocytherid ostracods with the de- scriptions of three new species.—Proceedings of the Biological Society of Washington 96(4):770-779. Ankylocythere carpenteri, Ascetocythere jezerinaci, Ornithocythere thomai. 1983f. Translation of Cambarinos de la Fauna Mexi- cana (Crustacea Decapoda) [Crayfishes of Mexico Crustacea: Decapoda], Alejandro Villalobos. Smith- 494 sonian Institution Libraries & National Science Foundation. 276 pp. Washington, D.C. 1984a. On the distribution of the crayfish genus Pro- cambarus (Decapoda: Cambaridae).—Journal of Crustacean Biology 4(1):12—24. 1984b. Burr, B. M., & H. H. Hobbs, Jr. Additions to the crayfish fauna of Kentucky, with new locality records for Cambarellus shufeldtii.—Transactions of the Kentucky Academy of Sciences 45(1—2):14—18. 1985a. Hobbs, H. H., Jr., & P H. Carlson. A new mem- ber of the genus Distocambarus (Decapoda: Cam- baridae) from the Saluda Basin, South Carolina.— Proceedings of the Biological Society of Washing- ton 98(1):81—89. D. youngineri. 1985b. Hobbs, H. H., Jr, & H. W. Robison. A new burrowing crayfish (Decapoda: Cambaridae) from southwestern Arkansas.—Proceedings of the Bio- logical Society of Washington 98(4):1035—1041. Fallicambarus harpi. 1986a. Andolshek, M. D., & H. H. Hobbs, Jr. The entocytherid ostracod fauna of southeastern Geor- gia.—Smithsonian Contributions to Zoology 424:43 pp. Ankylocythere spargosis, Entocythere prisma. 1986b. Hobbs, H. H., Jr, & R. Franz. New troglobitic crayfish with comments on its relationship to epi- gean and other hypogean crayfishes of Florida. Journal of Crustacean Biology 6(3):509-—519. Pro- cambarus delicatus. 1986c. Hobbs, H. H., Jr, & A. G. Grubbs. Notes on the crayfish Procambarus (Ortmannicus) xilitlae (Decapoda: Cambaridae).—Proceedings of the Bio- logical Society of Washington 99(4):735-738. 1986d. Highlights of a half century of crayfishing. Pp. 12—23 in P. Brinck, ed., Freshwater Crayfish VI, Pa- pers from the Sixth International Symposium of As- tacology, Lund Sweden, 13-15 August 1984. 281 pp. International Association of Astacology. 1986e. A new troglobitic crab (Crustacea: Decapoda: Pseudothelphusidae) from Belize. Pp. 1—4 in J. R. Reddell, ed., Studies on the cave and endogean fau- na of North America.—Texas Memorial Museum Speleological Monographs 1:1—4. Typhlopseudoth- elphusa acanthochela. 1987a. On the identity of Astacus (Cambarus) mexi- canus Erichson (1846) and Cambarus aztecus Saus- sure (1857) (Decapoda: Cambaridae) with the de- scription of Procambarus olmecorum, new species, from Veracruz, Mexico.—Proceedings of the Bio- logical Society of Washington 100(1):198—215. 1987b. A review of the crayfish genus Astacoides (De- capoda: Parastacidae).—Smithsonian Contributions to Zoology 443:50 pp. A. crosnieri, A. petiti. 1987c. Hobbs, H. H., Jr, & M. Whiteman. A new, economically important crayfish (Decapoda: Cam- baridae) from the Neches River Basin, Texas, with a key to the subgenus Fallicambarus.—Proceedings of the Biological Society of Washington 100(2): 403-411. Fallicambarus devastator. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1987d. Hobbs, H. H., Jr., & A. V. Brown. A new trog- lobitic crayfish from northwestern Arkansas (Decap- oda: Cambaridae).—Proceedings of the Biological Society of Washington 100(4):1040—1048. Camba- rus aculabrum. 1988a. Crayfish distribution, adaptive radiation and evolution. Pp. 52-82 in D. M. Holdich & R. S. Low- ery, eds., Freshwater crayfish: biology, management and exploitation. 498 pp. Croom Helm: London and Sydney; Timber Press: Portland, Oregon. 1988b. Hobbs, H. H., Jr., & H. W. Robison. The cray- fish subgenus Girardiella (Decapoda: Cambaridae) in Arkansas, with the descriptions of two new spe- cies and a key to the members of the Gracilis group in the genus Procambarus.—Proceedings of the Bi- ological Society of Washington 101(2):391—413. Procambarus ferrugineus, P. regalis. 1988c. Hobbs, H. H., Jr, & W. L. Pflieger. Cambarus (Erebicambarus) maculatus, new crayfish (Decapo- da: Cambaridae) from the Meramec River Basin of Missouri.—Proceedings of the Biological Society of Washington 101(3):644—-652. 1989a. Burrowing crayfishes in Virginia. The Virginia Explorer 5(2):5-7. 1989b. Hobbs, H. H., Jr., & D. J. Peters. New records of entocytherid ostracods infesting burrowing cray- fishes, with the description of a new species, Asce- tocythere stockeri.—Proceedings of the Biological Society of Washington 102(2):324—330. 1989c. Rodriguez, G., & H. H. Hobbs, Jr. Freshwater crabs associated with caves in southern Mexico and Belize, with descriptions of three new species (Crus- tacea: Decapoda).—Proceedings of the Biological Society of Washington 102(2):394—400. Odonto- thelphusa monodontis, Potamocarcinus leptomelus, Typhlopseudothelphusa hyba. 1989d. Rodriguez, G. & H. H. Hobbs, Jr. A new cav- ernicolous crab, Zilchia falcata, from Guatemala, with notes on the genera of the Potamocarcinini (Crustacea Decapoda, Pseudothelphusidae).—Bul- letin du Muséum National d’ Histoire Naturelle, Par- is, series 4, 11(section A) (1):183—192. 1989e. Hobbs, H. H. Jr., & H. H. Hobbs III. New lo- cality records for two poorly known Mexican fresh- water shrimps (Decapoda, Palaemonidae).—Crusta- ceana 57(2):220—222. 1989f. Hobbs, Jr., H. H., & H. W. Robison. On the crayfish genus Fallicambarus (Decapoda: Cambar- idae) in Arkansas, with notes on the Fodiens com- plex and descriptions of two new species.—Pro- ceedings of the Biological Society of Washington 102(3):651-697. F. gilpini, F. petilicarpus. 1989g. An illustrated checklist of the American cray- fishes (Decapoda: Astacidae, Cambaridae, and Par- astacidae).—Smithsonian Contributions to Zoology 480:236 pp. 1990a. On the crayfishes (Decapoda: Cambaridae) of the Neches River Basin of eastern Texas with the VOLUME 111, NUMBER 3 descriptions of three new species.—Proceedings of the Biological Society of Washington 103(3):573— 597. Procambarus kensleyi, P. nechesae, P. nigro- cinctus. 1990b. Hobbs, H. H., Jr., & H. H. Hobbs III. A new crayfish (Decapoda: Cambaridae) from southeastern Texas.—Proceedings of the Biological Society of Washington 103(3):608-613. Procambarus zonan- gulus. 1991a. Hobbs, H. H., Jr, & M. Whiteman. Notes on the burrows, behavior, and color of the crayfish Fal- licambarus (F.) devastator (Decapoda: Cambari- dae).—The Southwestern Naturalist 36(1):127—135. 1991b. Hobbs, H. H., Jr, & R. Franz. A new troglob- itic crayfish, Procambarus (Lonnbergius) morrisi, (Decapoda: Cambaridae) from Florida.—Proceed- ings of the Biological Society of Washington 104(1):55-63. 1991c. Hobbs, H. H., Jr, & D. J. Peters. Additional records of entocytherid ostracods infesting burrow- ing crayfishes, with the description of five new spe- cies.—Proceedings of the Biological Society of Washington 104(1):64—75. Ankylocythere prolata, Dactylocythere guyandottae, D. lepta, D. pygidion, Plectocythere kentuckiensis. 1991d. Hobbs, H. H., Jr, & H. H. Hobbs III. An il- lustrated key to the crayfishes of Florida (based on first form males).—Florida Scientist 54(1):13—24. 199le. Procambarus (Girardiella) steigmani, a new crayfish (Decapoda, Cambaridae) from a long-grass prairie in northeastern Texas.—Proceedings of the Biological Society of Washington 104(2):309-316. 1991f. A new generic assignment for a South Ameri- can crayfish (Decapoda: Parastacidae) with revised diagnoses of the South American genera and com- ments on the parastacid mandible.—Proceedings of the Biological Society of Washington 104(4):800— 811. Virilastacus. 1992. Hobbs, H. H., Jr., & R. Franz. Procambarus (Ortmannicus) attiguus, a new troglobitic crayfish (Decapoda, Cambaridae) from the Saint Johns River Basin, Florida.—Proceedings of the Biological So- ciety of Washington 105(2):359-—365. 1993. Hobbs, H. H., Jr, & D. J. Peters. New records of entocytherid ostracods infesting burrowing and cave-dwelling crayfishes, with descriptions of two new species.—Proceedings of the Biological Soci- ety of Washington 106(3):455—466. Dactylocythere cryptoteresis, Phymocythere lophota. 1994. Hobbs, H. H., Jr., & R. W. Bouchard. Cambarus (Cambarus) angularis, a new crayfish (Decapoda: Cambaridae) from the Tennessee River Basin of northeastern Tennessee and Virginia.—Jeffersoniana 5:1-13. 1995a. Hobbs, H. H., Jr., & H. H. Hobbs III. Macro- brachium catonium, a new troglobitic shrimp from the Cayo District of Belize (Crustacea: Decapoda: 495 Palaemonidae).—Proceedings of the Biological So- ciety of Washington 108(1):50-—53. 1995b. Hobbs, H. H., Jr., & H. H. Hobbs III. Procam- barus (Ortmannicus) nueces (Decapoda: Cambari- dae), a new crayfish from the Nueces River Basin, Texas.—Proceedings of the Biological Society of Washington 108(1):54—60. At the time of his death, Hobbs was com- pleting a manuscript entitled “‘A compara- tive study of functional morphology of the male reproductive systems in the Astacidea (Crustacea: Decapoda) with emphasis on the freshwater crayfishes.”’ Hobbs III plans to complete preparing this manuscript for publication. Taxa Named by Horton H. Hobbs, Jr. The family, genera and subgenera, spe- cies and subspecies named by Hobbs and colleagues are listed alphabetically and are cross-referenced to Hobbs’s bibliography, above. We provide the repository and cat- alogue number for all holotypes of species and subspecies. Most of the holotypes of taxa named by Hobbs are in the collections of the National Museum of Natural History, Smithsonian Institution, Washington (USNM). One holotype is in The Natural History Museum, London (BMNH) and several are in the Muséum National d’ Histoire Naturelle, Paris (MNHN) as well as The Academy of Natural Sciences of Philadelphia (ANSP). When a holotype has been deposited in a museum other than the USNM, catalogue numbers of USNM para- types are provided. Acucauda Hobbs, 1972a. Alpheopsis stygicola Hobbs, 1973d. Holo- type USNM 143629. Ankylocythere barbouri Villalobos Figu- eroa & Hobbs, 1974b. Holotype USNM 149159. Ankylocythere carpenteri Hobbs & Mc- Clure, 1983e. Holotype USNM 204402. Ankylocythere harmani Hobbs, 1966f. Ho- lotype USNM 123532. Ankylocythere hyba Hobbs & Walton, 1963a. Holotype USNM 108016. 496 Ankylocythere maya Hobbs, 1971b. Holo- type USNM 128822. Ankylocythere prolata Hobbs & Peters, 1991c. Holotype USNM 235511. Ankylocythere spargosis Andolshek & Hobbs, 1986. Holotype USNM 213651. Ankylocythere toltecae Hobbs, 1971b. Ho- lotype USNM 128823. Ankylocythere villalobosi Hobbs, Holotype USNM 128825. Aphelocythere Hobbs & Peters, 1977b. Aphelocythere acuta Hobbs & Peters, 1977b. Holotype USNM 155324. Ascetocythere batchi Hobbs & Walton, 1968d. Holotype USNM 123321. Ascetocythere bouchardi Hobbs & Walton, 1975b. Holotype USNM 150640. Ascetocythere coryphodes Hobbs & Hart, 1966d. Holotype USNM 113449. Ascetocythere cosmeta Hobbs & Hart, 1966d. Holotype USNM 113453. Ascetocythere didactylata Hobbs & Hart, 1966d. Holotype USNM 113448. Ascetocythere hoffmani Hobbs & Hart, 1966d. Holotype USNM 113441. Ascetocythere holti Hobbs & Walton, 1970b. Holotype USNM 126974. Ascetocythere hyperoche Hobbs & Hart, 1966d. Holotype USNM 113444. Ascetocythere jezerinaci Hobbs & Mc- Clure, 1983e. Holotype USNM 204400. Ascetocythere lita Hobbs & Hobbs III, 1970c. Holotype USNM 126251. Ascetocythere myxoides Hobbs & Hart, 1966d. Holotype USNM 113451. Ascetocythere ozalea Hobbs & Hart, 1966d. Holotype USNM 113442. Ascetocythere pseudolita Hobbs & Walton, 1975b. Holotype USNM 150642. Ascetocythere riopeli Hobbs & Walton, 1976d. Holotype USNM 155317. Ascetocythere sclera Hobbs & Hart, 1966d. Holotype USNM 113445. Ascetocythere stockeri Hobbs & Peters, 1989b. Holotype USNM 240114. Ascetocythere triangulata Hobbs & Walton, 1975b. Holotype USNM 150643. Ascetocythere veruta Hobbs & Walton, 1975b. Holotype USNM 150645. HO Well: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Astacoides crosnieri Hobbs, 1987b. Holo- type MNHN As328; paratype USNM NST. Astacoides petiti Hobbs, 1987b. Holotype MNHN As211; paratype USNM 218799. Atya brachyrhinus Hobbs & Hart, 1982e. Holotype BMNH 1972:539; paratype USNM 184857. Austrocambarus Hobbs, 1972a. Aviticambarus Hobbs, 1969c. Barbicambarus Hobbs, 1969c. Bouchardina Hobbs, 1977a. Bouchardina robisoni Hobbs, 1977a. Ho- lotype USNM 147146. Cambarellus blacki Hobbs, 1980b. Holo- type USNM 148901. Cambarellus chihuahuae Hobbs, Holotype USNM 148895. Cambarellus diminutus Hobbs, 1945c. Ho- lotype USNM 81554. Cambarellus ninae Hobbs, 1950a. Holotype USNM 89768. Cambarellus prolixus Villalobos-Figueroa & Hobbs, 198la. Holotype USNM 177206. Cambarellus puer Hobbs, 1945c. Holotype USNM 81556. Cambarellus schmitti Hobbs, 1942d. Ho- lotype USNM 81291. Cambaridae Hobbs, 1942d. Cambarus acanthura Hobbs, 1981b. Holo- type USNM 129758. Cambarus aculabrum Hobbs & Brown, 1987d. Holotype USNM 219149. Cambarus angularis Hobbs & Bouchard, 1994. Holotype USNM 260252. Cambarus blandingii cuevachicae Hobbs, 1941a. Holotype USNM 80030. Cambarus bouchardi Hobbs, 1970e. Holo- type USNM 130295. Cambarus brachydactylus Hobbs, 1953a. Holotype USNM 93155. Cambarus byersi Hobbs, 1941b. Holotype USNM 79342. Cambarus catagius Hobbs & Perkins, 1967c. Holotype USNM 117799. Cambarus chaugaensis Prins & Hobbs, 1972d. Holotype USNM 131926. 1980b. VOLUME 111, NUMBER 3 Cambarus conasaugaensis Hobbs & Hobbs III, 1962a. Holotype USNM 107156. Cambarus coosae Hobbs, 1981b. Holotype USNM 145603. Cambarus coosawattae Hobbs, 1981b. Ho- lotype USNM 148112. Cambarus cracens Bouchard & Hobbs, 1976b. Holotype USNM 146082. Cambarus cristatus Hobbs, 1955a. Holo- type USNM 96985. Cambarus cryptodytes Hobbs, 1941b. Ho- lotype USNM 79339. Cambarus cumberlandensis Hobbs & Bou- chard, 1973a. Holotype USNM 132989. Cambarus cymatilis Hobbs, 1970e. Holo- type USNM 129860. Cambarus englishi Hobbs & Hall, 1972f. Holotype USNM 131700. Cambarus fasciatus Hobbs, 1981b. Holo- type USNM 147917. Cambarus floridanus Hobbs, 1941b. Holo- type USNM 79341. Cambarus friaufi Hobbs, 1953a. Holotype USNM 93157. Cambarus gentryi Hobbs, 1970d. Holotype USNM 130283. Cambarus georgiae Hobbs, 1981b. Holo- type USNM 118944. Cambarus halli Hobbs, 1968a. Holotype USNM 129288. Cambarus harti Hobbs, 1981b. Holotype USNM 148348. Cambarus hedgpethi Hobbs, 1948f. Holo- type USNM 85146. Cambarus hiwasseensis Hobbs, 1981b. Ho- lotype USNM 129366. Cambarus howardi Hobbs & Hall, 1969d. Holotype USNM 129866. Cambarus hubrichti Hobbs, 1952d. Holo- type USNM 92295. Cambarus hubbelli Hobbs, 1940a. Holo- type USNM 76593. Cambarus jonesi Hobbs & Barr, 1960b. Holotype USNM 104407. Cambarus kilbyi Hobbs, 1940a. Holotype USNM 76594. Cambarus lucifugus alachua Hobbs, 1940a. Holotype USNM 76592. 497 Cambarus lucifugus lucifugus Hobbs, 1940a. Holotype USNM 77916. Cambarus lunzi Hobbs, 1940b. Holotype USNM 79301. Cambarus maculatus Hobbs & Pflieger, 1988c. Holotype USNM 219292. Cambarus manningi Hobbs, 1981b. Holo- type USNM 147911. Cambarus nerterius Hobbs, 1964c. Holo- type USNM 111295. Cambarus nodosus Bouchard & Hobbs, 1976b. Holotype USNM 146756. Cambarus obeyensis Hobbs & Shoup, 1947b. Holotype USNM 82260. Cambarus pallidus Hobbs, 1940a. Holotype USNM 76591. Cambarus parrishi Hobbs, 1981b. Holo- type USNM 144957. Cambarus parvoculus Hobbs & Shoup, 1947b. Holotype USNM 82259. Cambarus pictus Hobbs, 1940a. Holotype USNM 76596. Cambarus pristinus Hobbs, 1965b. Holo- type USNM 115528. Cambarus prominens Hobbs, 1966b. Ho- lotype USNM 115603. Cambarus rathbunae Hobbs, 1940a. Holo- type USNM 76595. Cambarus reduncus Hobbs, 1956a. Holo- type USNM 99183. Cambarus reflexus Hobbs, 1981b. Holotype USNM 148116. Cambarus rogersi Hobbs, 1938b. Holotype USNM 75120. Cambarus scotti Hobbs, 1981b. Holotype USNM 146479. Cambarus speciosus Hobbs, 1981b. Holo- type USNM 146023. Cambarus sphenoides Hobbs, 1968a. Ho- lotype USNM 129325. Cambarus spicatus Hobbs, 1956b. Holo- type USNM 99323. Cambarus strigosus Hobbs, 1981b. Holo- type USNM 148284. Cambarus tartarus Hobbs & Cooper, 1972e. Holotype USNM 131951. Cambarus truncatus Hobbs, 1981b. Holo- type USNM 116966. 498 Cambarus unestami Hobbs & Hall, 1969d. Holotype USNM 129863. Cambarus zophonastes Hobbs & Bedinger, 1964b. Holotype USNM 108356. Capillicambarus Hobbs, 1972a. Creaserinus Hobbs, 1973e. Cymocythere gonia Hobbs & Hart, 1966d. Holotype USNM 113455. Dactylocythere apheles Hobbs & Walton, 1976d. Holotype USNM 155319. Dactylocythere astraphes Hobbs & Walton, 1977f. Holotype USNM 169073. Dactylocythere brachydactylus Hobbs & Walton, 1976d. Holotype USNM 155321. Dactylocythere brachystrix Hobbs & Wal- ton, 1966e. Holotype USNM 111251. Dactylocythere coloholca Hobbs & Hobbs III, 1970c. Holotype USNM 126253. Dactylocythere cooperorum Hobbs & Wal- ton, 1968d. Holotype USNM 123322. Dactylocythere corvus Hobbs & Walton, 1977f. Holotype USNM 169075. Dactylocythere crena Hobbs & Walton, 1975b. Holotype USNM 150646. Dactylocythere cryptoteresis Hobbs & Pe- ters, 1993. Holotype USNM 260073. Dactylocythere demissa Hobbs & Walton, 1976d. Holotype USNM 155323. Dactylocythere enoploholea Hobbs & Wal- ton, 1970b. Holotype USNM 126973. Dactylocythere guyandottae Hobbs & Pe- ters, 1991c. Holotype USNM 235513. Dactylocythere isabelae Hobbs & Peters, 1977b. Holotype USNM 155326. Dactylocythere jeanae Hobbs, 1967a. Ho- lotype USNM 113475. Dactylocythere lepta Hobbs & Peters, 1991c. Holotype USNM 235514. Dactylocythere macroholca Hobbs & Hobbs III, 1970c. Holotype USNM 126255. Dactylocythere myura Hobbs & Walton, 1970b. Holotype USNM 126975. Dactylocythere pachysphyrata Hobbs & Walton, 1966e. Holotype USNM 111253. Dactylocythere peedeensis Hobbs & Peters, 1977b. Holotype USNM 155328. Dactylocythere phoxa Hobbs, 1967a. Ho- lotype USNM 113477. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Dactylocythere prinsi Hobbs & Walton, 1968d. Holotype USNM 123325. Dactylocythere prominula Hobbs & Wal- ton, 1977f. Holotype USNM 169077. Dactylocythere pughae Hobbs & Hobbs III, 1970c. Holotype USNM 126257. Dactylocythere pygidion Hobbs & Peters, 1991c. Holotype USNM 235516. Dactylocythere scissura Hobbs & Walton, 1975b. Holotype USNM 150648. Dactylocythere spinata Hobbs & Walton, 1970b. Holotype USNM 126972. Dactylocythere spinescens Hobbs & Wal- ton, 1977f. Holotype USNM 169079. Dactylocythere xystroides Hobbs & Walton, 1963a. Holotype USNM 108029. Depressicambarus Hobbs, 1969c. Distocambarus Hobbs, 1981b. Distocambarus carlsoni Hobbs, 1983d. Ho- lotype USNM 178599. Distocambarus crockeri Hobbs & Carlson, 1983c. Holotype USNM 178582. Distocambarus youngineri Hobbs & Carl- son, 1985a. Holotype USNM 208414. Donnaldsoncythere ardis Hobbs & Walton, 1963b. Holotype USNM 108018. Donnaldsoncythere cayugaensis Hobbs & Walton, 1966e. Holotype USNM 111255. Donnaldsoncythere ileata Hobbs & Walton, 1963b. Holotype USNM 108027. Donnaldsoncythere leptodrilus Hobbs & Peters, 1977b. Holotype USNM 155330. Donnaldsoncythere scalis Hobbs & Walton, 1963b. Holotype USNM 108024. Donnaldsoncythere truncata Hobbs & Wal- ton, 1963b. Holotype USNM 108021. Entocythere ambophora Walton & Hobbs, 1959c. Holotype USNM 105953. Entocythere arcuata Hart & Hobbs, 1961b. Holotype ANSP 6233. Entocythere asceta Hobbs & Walton, 1962b. Holotype USNM 108174. Entocythere barri Hart & Hobbs, 1961b. Holotype ANSP 6231. Entocythere chalaza Hobbs & Walton, 1962b. Holotype USNM 108179. Entocythere costata Hobbs & Peters, 1977b. Holotype USNM 155334. VOLUME 111, NUMBER 3 Entocythere cyma Hobbs & Walton, 1960a. Holotype USNM 105962. Entocythere daphnioides Hobbs, 1955b. Holotype USNM 98406. Entocythere falcata Hobbs & Walton, 1961la. Holotype USNM 105966. Entocythere hamata Hobbs, 1957a. Holo- type USNM 100938. Entocythere hiwasseensis Hobbs & Walton, 1961a. Holotype USNM 105963. Entocythere kanawhaensis Hobbs & Wal- ton, 1966e. Holotype USNM 111257. Entocythere lucifuga Walton & Hobbs, 1959c. Holotype USNM 105957. Entocythere mecoscapha Hobbs & Walton, 1960a. Holotype USNM 105960. Entocythere pholetera Hart & Hobbs, 1961b. Holotype ANSP 6238. Entocythere phyma Hobbs & Walton, 1962b. Holotype USNM 108178. Entocythere prionata Hart & Hobbs, 1961b. Holotype ANSP 6235. Entocythere prisma Andolshek & Hobbs, 1986a. Holotype USNM 213649. Entocythere reddelli Hobbs & Walton, 1968d. Holotype USNM 123330. Entocythere runki Hobbs, 1955b. Holotype USNM 98410. Entocythere simondsi Hobbs & Walton, 1960a. Holotype USNM 105956. Entocythere steevesi Hart & Hobbs, 1961b. Holotype ANSP 6236. | Entocythere tuberosa Hart & Hobbs, 1961b. Holotype ANSP 6340. Entocythere tyttha Hobbs & Hobbs III, 1970c. Holotype USNM 126258. Entocythere ungulata Hart & Hobbs, 1961b. Holotype ANSP 6234. Entocythere xania Hart & Hobbs, 1961b. Holotype ANSP 6237. Erebicambarus Hobbs, 1969a. Exilicambarus Bouchard & Hobbs, 1976d. Fallicambarus Hobbs, 1969c. Fallicambarus caesius Hobbs, 1975c. Ho- lotype USNM 144921. Fallicambarus danielae Hobbs, 1975c. Ho- lotype USNM 145997. Fallicambarus devastator Hobbs & White- man, 1987c. Holotype USNM 218546. 499 Fallicambarus gilpini Hobbs & Robison, 1989f. Holotype USNM 219511. Fallicambarus harpi Hobbs & Robison, 1985b. Holotype USNM 217946. Fallicambarus hortoni Hobbs & Fitzpat- rick, 1970a. Holotype USNM 129895. Fallicambarus jeanae Hobbs, 1973e. Ho- lotype USNM 144672. Fallicambarus petilicarpus Hobbs & Rob- ison, 1989f. Holotype USNM 219507. Fallicambarus spectrum Hobbs, 1973e. Ho- lotype USNM 144674. Fitzcambarus Hobbs, 1983d. Geocythere nessoides Hobbs & Hobbs III, 1970c. Holotype USNM 126259. Hagenides Hobbs, 1972a. Harpagocythere baileyi Hobbs & Peters, 1977b. Holotype USNM 155332. Harpagocythere tertius Hobbs & Walton, 1968d. Holotype USNM 123329. Hiaticambarus Hobbs, 1969c. Jugicambarus Hobbs, 1969c. Lacunicambarus Hobbs, 1969c. Leconticambarus Hobbs, 1972a. Litocythere Hobbs & Walton, 1968d. Litocythere lucileae Hobbs & Walton, 1968d. Holotype USNM 123327. Lonnbergius Hobbs, 1972a. Lordocythere Hobbs & Hobbs III, 1970c. Lordocythere petersi Hobbs & Hobbs III, 1970c. Holotype USNM 126262. Macrobrachium catonium Hobbs & Hobbs III, 1995a. Holotype USNM 260328. Macrobrachium villalobosi Hobbs, 1973d. Holotype USNM 143633. Mexicambarus Hobbs, 1972a. Neopalaemon Hobbs, 1973b. Neopalaemon nahuatlus Hobbs, 1973b. Holotype USNM 143120. Odontothelphusa monodontis Rodriguez & Hobbs, 1989c. Holotype USNM 230078. Orconectes chickasawae Cooper & Hobbs, 1980c. Holotype USNM 144926. Orconectes cooperi Cooper & Hobbs, 1980c. Holotype USNM 147722. Orconectes holti Cooper & Hobbs, 1980c. Holotype USNM 147149. Orconectes incomptus Hobbs & Barr, 1972b. Holotype USNM 130299. 500 Orconectes leptogonopodus Hobbs, 1948b. Holotype USNM 82262. Orconectes marchandi Hobbs, 1948b. Ho- lotype USNM 82072. Orconectes propinquus erismophorous Hobbs & Fitzpatrick, 1962d. Holotype USNM 107597. Orconectes rhoadesi Hobbs, 1949b. Holo- type USNM 87953. Orconectes shoupi Hobbs, 1948a. Holotype USNM 84072. Orconectes virginiensis Hobbs, 1951a. Ho- lotype USNM 91659. Orconectes wrighti Hobbs, 1948e. Holo- type USNM 85144. Ornithocythere Hobbs, 1967a. Ornithocythere thomai Hobbs & McClure, 1983e. Holotype USNM 204405. Ornithocythere waltonae Hobbs, 1967a. Holotype USNM 113472. Palaemonetes lindsayi Villalobos Figueroa & Hobbs, 1974b. Holotype USNM 149161. Pennides Hobbs, 1972a. Phymocythere Hobbs & Hart, 1966d. Phymocythere lophota Hobbs & Peters, 1993. Holotype USNM 260074. Plectocythere johnsonae Hobbs & Hart, 1966d. Holotype USNM 113454. Plectocythere kentuckiensis Hobbs & Pe- ters, 1991c. Holotype USNM 235517. Potamocarcinus leptomelus Rodriguez & Hobbs, 1989c. Holotype USNM 230080. Procambarus ancylus Hobbs, 1958b. Ho- lotype USNM 115050. Procambarus apalachicolae Hobbs, 1942d. Holotype USNM 81272. Procambarus attiguus Hobbs & Franz, 1992. Holotype USNM 220683. Procambarus bivittatus Hobbs, 1942d. Ho- lotype USNM 81274. Procambarus capillatus Hobbs, 1971c. Ho- lotype USNM 131454. Procambarus caritus Hobbs, 1981b. Holo- type USNM 117598. Procambarus chacei Hobbs, 1958c. Holo- type USNM 101289. Procambarus clemmeri Hobbs, 1975c. Ho- lotype USNM 145607. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Procambarus cuetzalanae Hobbs, 1982a. Holotype USNM 177202. Procambarus delicatus Hobbs & Franz, 1986b. Holotype USNM 218528. Procambarus devexus Hobbs, 1981b. Ho- lotype USNM 148569. Procambarus echinatus Hobbs, 1956c. Ho- lotype USNM 99180. Procambarus econfinae Hobbs, 1942d. Ho- lotype USNM 81276. Procambarus elegans Hobbs, 1969e. Ho- lotype USNM 129892. Procambarus enoplosternum Hobbs, 1947a. Holotype USNM 82263. Procambarus epicyrtus Hobbs, 1958c. Ho- lotype USNM 101286. Procambarus escambiensis Hobbs, 1942d. Holotype USNM 81278. Procambarus ferrugineus Hobbs & Robi- son, 1988b. Holotype USNM 218841. Procambarus fitzpatricki Hobbs, 1971a. Holotype USNM 131205. Procambarus franzi Hobbs & Lee, 1976c. Holotype USNM 146992. Procambarus geminus Hobbs, 1975c. Ho- lotype USNM 145756. Procambarus geodytes Hobbs, 1942d. Ho- lotype USNM 82263. Procambarus gibbus Hobbs, 1969e. Holo- type USNM 129804. Procambarus hirsutus Hobbs, 1958b. Ho- lotype USNM 101148. Procambarus horsti Hobbs & Means, 1972c. Holotype USNM 132043. Procambarus howellae Hobbs, 1952c. Ho- lotype USNM 93158. Procambarus hybus Hobbs & Walton, 1957b. Holotype USNM 99581. Procambarus jaculus Hobbs & Walton, 1957b. Holotype USNM 99584. Procambarus kensleyi Hobbs, 1990a. Ho- lotype USNM 219772. Procambarus latipleurum Hobbs, 1942d. Holotype USNM 81281. Procambarus leitheuseri Franz & Hobbs, 1983a. Holotype USNM 178361. Procambarus leonensis Hobbs, 1942d. Ho- lotype USNM 81091. VOLUME 111, NUMBER 3 Procambarus lepidodactylus Hobbs, 1947c. Holotype USNM 84198. Procambarus lewisi Hobbs & Walton, 1959a. Holotype USNM 102467. Procambarus litosternum Hobbs, Holotype USNM 82261. Procambarus lophotus Hobbs & Walton, 1960c. Holotype USNM 104404. Procambarus lylei Fitzpatrick & Hobbs, 1971d. Holotype USNM 131533. Procambarus mancus Hobbs & Walton, 1957b. Holotype USNM 99578. Procambarus marthae Hobbs, 1975c. Ho- lotype USNM 145994. Procambarus medialis Hobbs, 1975c. Ho- lotype USNM 144942. Procambarus milleri Hobbs, 1971f. Holo- type USNM 131257. Procambarus morrisi Hobbs & Franz, 1991b. Holotype USNM 220374. Procambarus nechesae Hobbs, 1990a. Ho- lotype USNM 219733. Procambarus nigrocinctus Hobbs, 1990a. Holotype USNM 219426. Procambarus niveus Hobbs & Villalobos, 1964a. Holotype USNM 109076. Procambarus nueces Hobbs & Hobbs III, 1995b. Holotype USNM 260326. Procambarus oaxacae oaxacae Hobbs, 1973b. Holotype USNM 144341. Procambarus oaxacae reddelli Hobbs, 1973b. Holotype USNM 144346. Procambarus okaloosae Hobbs, 1942d. Ho- lotype USNM 81282. Procambarus olmecorum Hobbs, Holotype USNM 217626. Procambarus orcinus Hobbs & Means, 1972c. Holotype USNM 132031. Procambarus parasimulans Hobbs & Rob- ison, 1982d. Holotype USNM 177698. Procambarus pearsei plumimanus Hobbs & Walton, 1958d. Holotype USNM 100940. Procambarus pecki Hobbs, 1967e. Holo- type USNM 117684. Procambarus penni Hobbs, 1951b. Holo- type USNM 91662. Procambarus petersi Hobbs, 1981b. Holo- type USNM 144960. 1947a. 1987a. 501 Procambarus pubischelae Hobbs, 1942d. Holotype USNM 81284. Procambarus pubischelae deficiens Hobbs, 1981b. Holotype USNM 133315. Procambarus pycnogonopodus Hobbs, 1942d. Holotype USNM 81092. Procambarus pygmaeus Hobbs, 1942d. Ho- lotype USNM 81285. Procambarus raneyi Hobbs, 1953e. Holo- type USNM 95124. Procambarus regalis Hobbs & Robison, 1988b. Holotype USNM 219244. Procambarus reimeri Hobbs, 1979b. Ho- lotype USNM 148880. Procambarus roberti Villalobos Figueroa & Hobbs, 1974b. Holotype USNM 132345. Procambarus rodriguezi Hobbs, 1943c. Holotype USNM 81302. Procambarus rogersi campestris Hobbs, 1942d. Holotype USNM 79384. Procambarus rogersi expletus Hobbs & Hart, 1959b. Holotype USNM 102125. Procambarus rogersi ochlocknensis Hobbs, 1942d. Holotype USNM 79383. Procambarus sbordonii Hobbs, 1978. Ho- lotype USNM 146588. Procambarus seminolae Hobbs, 1942d. Ho- lotype USNM 81286. Procambarus shermani Hobbs, 1942d. Ho- lotype USNM 81287. Procambarus steigmani Hobbs, 1991e. Ho- lotype USNM 220525. Procambarus strenthi Hobbs, 1977e. Ho- lotype USNM 147725. Procambarus suttkusi Hobbs, 1953d. Ho- lotype USNM 93730. Procambarus talpoides Hobbs, 1981b. Ho- lotype USNM 115002. Procambarus tenuis Hobbs, 1950b. Holo- type USNM 90390. Procambarus texanus Hobbs, 1971c. Ho- lotype USNM 131457. Procambarus toltecae Hobbs, 1943c. Ho- lotype USNM 81301. Procambarus truculentus Hobbs, Holotype USNM 95670. Procambarus verrucosus Hobbs, 1952a. Holotype USNM 118675. 1954b. 502 Procambarus villalobosi Hobbs, 1969b. Holotype USNM 118644. Procambarus xilitlae Hobbs & Grubbs, 1982b. Holotype USNM 177140. Procambarus xochitlanae Hobbs, Holotype USNM 145610. Procambarus youngi Hobbs, 1942d. Holo- type USNM 81289. Procambarus zonangulus Hobbs & Hobbs III, 1990b. Holotype USNM 220297. Pseudothelphusa seiferti Hobbs, 1980d. Holotype USNM 173884. Psittocythere Hobbs & Walton, 1975b. Psittocythere psitta Hobbs & Walton, 1975b. Holotype USNM 150650. Puncticambarus Hobbs, 1969c. Remoticambarus Hobbs, 1972a. Scapulicambarus Hobbs, 1972a. Sphaeromicola coahuiltecae Hobbs & Hobbs, III, 1973c. Holotype USNM 143122. Tenuicambarus Hobbs, 1972a. Thermastrocythere Hobbs 1966e. Thermastrocythere harti Hobbs & Walton, 1966e. Holotype USNM 111258. Troglocambarus Hobbs, 1942b. Troglocambarus maclanei Hobbs, 1942b. Holotype USNM 79385. Typhlatya campecheae Hobbs III & Hobbs, 1976e. Holotype USNM 151903. Typhlatya mitchelli Hobbs II & Hobbs, 1976e. Holotype USNM 151904. Typhlopseudothelphusa acanthochela Hobbs, 1986e. Holotype USNM 216239. Typhlopseudothelphusa hyba Rodriguez & Hobbs, 1989c. Holotype USNM 150633. Uncinocythere stubbsi Hobbs & Walton, 1966e. Holotype USNM 111260. Uncinocythere warreni Hobbs & Walton, 1968d. Holotype USNM 123324. Uncinocythere zancla Hobbs & Walton, 1963a. Holotype USNM 108033. Uncinocythere zaruri Hobbs, 1971b. Holo- type USNM 128827. Veticambarus Hobbs, 1969c. Villalobosus Hobbs, 1972a. Virilastacus Hobbs, 1991f. Waltoncythere Hobbs & Peters, 1979a. 1975c. & Walton, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Zilchia falcata Rodriguez & Hobbs, 1989d. Holotype MNHN B20118. Acknowledgments Many of Hobbs’s colleagues, friends, and family contributed to this biography, and we are indebted to all of them for taking time to read a late draft and their interest, support, corrections, suggestions, and pho- tographs, all of which materially improved our effort. We acknowledge with thanks the following individuals: Fenner A. Chace, Jr., National Museum of Natural History, Smithsonian Institution, Washington, D.C.; John E. Cooper, North Carolina State Mu- seum of Natural Sciences, Raleigh; Joseph FE Fitzpatrick, Jr., University of South Ala- bama, Mobile; Richard Franz, Florida State Museum, University of Florida, Gaines- ville; Horton H. Hobbs, III, Wittenberg University, Springfield, Ohio; Brian Ken- sley and Elizabeth Nelson, National Muse- um of Natural History, Smithsonian Insti- tution, Washington, D.C.; James E Payne, Memphis State University, Tennessee; Dan- iel J. Peters, New Horizons Governor’s School for Science and Technology, Hamp- ton, Virginia; Rudolph Prins, Western Ken- tucky University, Bowling Green; Jean Pugh, Yorktown, Virginia; and Nina Hobbs Singleton, Philadelphia, Pennsylvania. Literature Cited All papers by Hobbs are cited above in his bibliography. They are not repeated here if cited in the text. Chace, Fenner A., Jr. 1951. The number of species of decapod and stomatopod Crustacea.—Journal of the Washington Academy of Sciences 41(11):370-372. Cooper, J. E., & M. R. Cooper. 1997. A new species of troglobitic crayfish of the genus Cambarus, subgenus Aviticambarus (Decapoda: Cambari- dae), endemic to White Spring Cave, Ala- bama.—Proceedings of the Biological Society of Washington 110(4):608-616. Davidson, A. 1979. North Atlantic Seafood. New York, The Viking Press, 512 pp. Dobbin, C. N. 1941. Fresh-water Ostracoda from Washington and other western localities.—Uni- VOLUME 111, NUMBER 3 versity of Washington Publications in Biology 4:174-246, 14 pls. Fitzpatrick, J. E, Jr. 1995. Horton H. Hobs, Jr. (29 March 1914-22 March 1994).—Journal of Crustacean Biology 15(4):797—799. . 1996. Horton H. Hobbs [sic], Jr. (29 March 1914-22 March 1994).—Crustaceana 69(2): 251-264. . 1995. Tribute to Horton H. Hobbs, Jr (29 March 1914—22 March 1994). Pp. ix, x in M. C. Geddes, D. R. Fielder, & A. M. M. Richard- son, eds., Freshwater crayfish X. Proceedings of the International Association of Astacology Tenth Symposium 1994, Adelaide, Australia, 637 pp. Hart, C. W., Jr. 1960. A new ostracod of the genus Entocythere from Pennsylvania.—Notulae Na- turae, Academy of Natural Sciences of Phila- delphia 335:4 pp. 503 Hoff, C. C. 1943. Two new ostracods of the genus Entocythere and records of previously described species.—Journal of the Washington Academy of Sciences 33(9):276—286. Hoffman, R. L. 1994. +Horton Holcombe Hobbs II (1914—1994).—Banisteria 4:36—-38. Klie, W. 1931. Crustacés Ostracodes. Campagne Speo- logique de C. Bolivar et R. Jeannel dans l’Amérique du Nord (1928), 3.—Biospeleolo- gica: Archives de Zoologie Expérimentale et Générale 71(3):333-—344. Rioja, E. 1943. Polimorfismo femenino en los ostra- codos del género Entocythere.—Anales del In- stituto de Biologia, Universidad Autonoma de México 14(2):567—585. Tressler, W. L. 1947. A check list of the known species of North American freshwater ostracods.—The American Midland Naturalist 38:698—707. Wolf, B. 1934-1938. Animalium Cavernarum Catalo- gus. 2 volumes, s’Gravenhage: W. Junk. 918 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):504—510. 1998. Franconictis (Mammalia: Carnivora) from the Late Oligocene of eastern Kazakstan Spencer G. Lucas, Robert J. Emry, and Pyruza A. Tleuberdina (SGL) New Mexico Museum of Natural History and Science, 1801 Mountain Road N.W., Albuquerque, New Mexico 87104, U.S.A.; (RJE) Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (PAT) Institute of Zoology, National Academy of Science, Akademgorodok, Almaty 480032, Kazakstan Abstract.—The right dentary of a small carnivore from near Ayaguz in east- ern Kazakstan is identified as Franconictis sp. aff. F. vireti (Dehm). Francon- ictis has previously been reported only from the early Miocene of Europe. Previous reports of Plesictis from Asia are not well founded. Franconictis from Ayaguz extends the geographic range of the genus into Asia and its temporal range back to the late Oligocene. Fossil mammals from eastern Kazakstan are best known from the richly fossiliferous Paleogene-Neogene strata of the Zaysan ba- sin (Fig. 1). Some Paleogene and more ex- tensive Neogene mammalian assemblages are also known from the intermontane ba- sins of the northern Tien Shan drained by the Ily River and its tributaries (Tleuberdina et al. 1993, Lucas & Bayshashov 1996, Emry et al. 1997). However, between the Ily and Zaysan basins, relatively few Ce- nozoic fossil mammals are known, in part because outcrops are limited in the relative- ly low topography of the Balkash-Alakol drainage basin. Here, we add to this sparse record a jaw of the mustelid carnivore Franconictis, collected near Ayaguz. Fran- conictis has previously been known only from the early Miocene of Europe (Wolsan 1993, Morlo 1996); this specimen is the first record of the genus from Asia and ex- tends its temporal range back to the late Oligocene. Abbreviations used.—When used in den- tal notations, upper case letters denote up- per (skull) teeth and lower case letters de- note lower (dentary) teeth. Institutional ab- breviations used are: AMNH—American Museum of Natural History, New York; KAN-Kazak Academy of Sciences, Alma- ty. Systematic Paleontology Order Carnivora Bowdich, 1821 Family Mustelidae Fischer von Waldheim, 1817 Genus Franconictis Wolsan, 1993 Franconictis sp. aff. F. vireti (Dehm 1950) Fig. 2 Referred specimen.—KAN 401-67, hor- izontal ramus of right dentary with pl—ml, from lower Miocene strata termed “Aral svita’’ near Ayaguz, Kazakstan. Description.—The dentary is narrow and shallow with two mental foramina on its la- bial aspect—one under the anterior root of the p2 and the other under the center of the p3. The ascending ramus is broken, but ap- pears to have been tall and thin, with the anterior border of the coronoid fossa sharp- ly defined. There is a dorso-ventrally oval alveolus for the canine at the anterior edge of the dentary as preserved. Evidently the postca- nine diastema was either short or nonexis- tent. The pl is a small, well worn trenchant tooth with a single root. The crown consists VOLUME 111, NUMBER 3 Miles 0 300 ———————— 400 200 300 400 500 Kilometers 0 —_—" NS < N (Gad 5 ot, J Worn ~~ e vv Aktyubinsk Kazakstan _ Caspian STEAK = : > A ’ | we . ’ Sea NN s | Uzbekistan ¢ \ Turkmenistan ‘\ es a Afghanistan Fig. 1. Ily basin in the eastern part of the country. of an anteriorly situated main cusp (proto- conid) with a long posterior surface sloping to a low heel. The p! length = 2.4 mm, width = 1.3 mm. The p2 has two distinct roots and a trenchant crown dominated by the tall pro- toconid. There is a very small anterior cin- gulid and a low, broad heel defined by the lingual cingulid which continues around the posterior margin of the tooth. The p2 length = 3.3 mm, width = 1.6 mm. The p3 is similar to the p2 but larger and @ Kyzyl-Orda “1 Tashkent 47” ace cf NaS ww F usd 3 P08) (Gose pt cence 1 505 Novosibirsk Heese eres, gis .7*—Petropaviovskt- Nu, wie Semipalatinsk e Zaysan Ayaguz~_ a) Alma-Ata : 0 oC | My, 4 na Kyrgyzstan _ ) ( g ~ uN 2 \ Tajikistan y Cut : SF: Alpen Gee en ee » ) Cae d barca Gc Aime a Pakistan R Map of Kazakstan showing location of the Zaysan basin, the Ayaguz fossil mammal locality and the has a relatively longer anterior slope. A very slight swelling low on the posterior slope of the protoconid is an incipient post- protoconid cuspid. The p4 has a similar overall shape, except the anterior cingulid bears a distinct cuspid, there is a small cus- pid (?metaconid) on the posterior slope of the protoconid, and the posterolingual cin- gulid is a sharp ridge produced into a small posterior accessory cuspid. The p3 length = 3.8 mm, width = 1.8 mm; p4 length = 4.6 mm, width = 2.1 mm. 506 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. m1, occlusal (A), lingual (B) and labial (C) views. Bar scale = 5 mm. The m1 is the only molar tooth remain- ing in the jaw, and it is substantially larger than the other teeth. The large, postero-la- bially canted protoconid dominates the tri- gonid. A relatively small and low metaco- nid is directly lingual to the protoconid, and a larger but lower paraconid projects an- tero-lingually, so that the trigonid basin is broadly open lingually. A carnassial notch interrupts the paracristid. The low talonid consists of a basin surrounded by a rela- tively sharp rim, which is much lower lin- gually than labially. The hypoconid is rel- atively large and the entoconid is present but poorly differentiated as a small cuspid on the posterolingual rim of the talonid; be- tween the hypoconid and entoconid are two small, bead-like cuspids. The m1 length = 6.4 mm, width = 2.9 mm. Franconictis sp. aff. F. vireti (Dehm) from eastern Kazakstan, KAN 401-67, right dentary with pl— The m2 is missing, but its position is in- dicated by two alveoli, the posterior of which is well up on the slope of the as- cending ramus. It was obviously a much smaller tooth than the ml. Identification.—The presence of a single- rooted pl supports assignment of KAN 401-67 to the Mustelidae sensu Wolsan (1993). Among mustelids, its closest simi- larity is to “‘Plesictis,” particularly the rel- atively small species “‘P.”’ vireti from Win- tershof-West, Germany (e.g., Dehm 1950, figs. 92-99). Points of close resemblance include the shallow dentary, single-rooted pl, lack of a posterior cuspid on p2 or p3, m1 trigonid less than three times as long as the talonid, m1 protoconid relatively low, broad and posteriorly inclined, ml meta- conid higher than paraconid, m1 entoconid VOLUME 111, NUMBER 3 and entoconulid poorly differentiated, m2 two-rooted, and m2 alveoli on the ascend- ing ramus. In a re-evaluation of the phylogeny and classification of European mustelids, Wol- san (1993) recently removed Plesictis vireti from Plesictis and placed it in the new ge- nus Franconictis. Given its strong similar- ity to P. vireti, we identify KAN 401-67 as Franconictis, but do not attempt a definite species-level identification of so incomplete a specimen. Therefore, we identify KAN 401-67 as Franconictis sp. aff. F. vireti. M. Wolsan (in litt., 1997) notes that KAN 401- 67 is slightly smaller and “‘more primitive”’ than F. vireti and probably represents a new species of Franconictis, but we believe the specimen is inadequate material upon which to base a new species. Plesictis in Asia To our knowledge (also see Russell & Zhai 1987, Werdelin 1996), Plesictis has been reported only twice from Asia. Ga- buniya (1964) first reported Plesictis based on an isolated canine, fragment of a distal humerus and partial metacarpal (Gabuniya 1964, figs. 17-19) from the Benara locality (late Oligocene) in western Georgia. These fossils represent a small carnivore, but are not sufficient to identify Plesictis. There- fore, we identify them as Carnivora, inde- terminate. Mellett (1968) reported cf. Plesictis sp. from the upper part of the Hsanda Gol For- mation (Mongolia, late Oligocene) based on AMNH 21654, a right dentary with m1—2 (Fig. 3). In this specimen, m1 is much larg- er than m2 and has a long, tall trigonid with the basin broadly open lingually. Its proto- conid is large, tall and inclined postero-la- bially, and the metaconid is a much lower, pointed cuspid lingual to the protoconid. The paraconid is intermediate in size be- tween the protoconid and metaconid and is massive and projects anteriorly. A carnas- sial notch interrupts the paracristid. The tal- onid is very low and cup-like with a distinct 507 Fig. 3. AMNH 21654, Stenoplesictis constans from the Hsanda Gol Formation of Mongolia, right dentary fragment with m1—2, lingual (A) and occlusal (B) views. Bar scale = 5 mm. hypoconid. The m1 length = 8.4 mm, width = 4.0 mm. The m2 is a small, oval tooth with a three-cusped trigonid and a low talonid with a shelf-like posterior rim. The m2 length = 3.6 mm, width = 2.8 mm. This specimen conforms well to Steno- plesictis, particularly in its relatively tall m1 trigonid, prevallid shear on ml, small m2 with a well developed trigonid and narrow, trenchant talonid, and lack of m3 (Dashzev- eg 1996). Indeed, AMNH 21654 closely re- sembles Stenoplesictis constans from the Hsanda Gol Formation (Matthew & Grang- er 1924, fig. 6F), to which we assign it. Therefore, there are no well founded reports of Plesictis in Asia. Biostratigraphy and Biochronology The locality from which KAN 401-67 was collected is on the right bank of the Ayaguz River, 29 km downstream from the town of Ayaguz, in eastern Kazakstan. The vertebrate-producing strata (Fig. 4) here rest with erosional unconformity on red beds 508 allUViUI ie cement ae Khorgos 2.22.02 2°%,0202%,5 svita Pavlodar legend svita — conglomerate ee sandstone ~—SS [] mudstone So Sa Se [__] limestone a) ‘ = metamorphic —— aS Aral Pots svita ae | Avagtz 6:00:60. ¢ gam) <= fossil pee locality ae Seeee = a Paleogene ->—————4] — 1 on u S woes we ! SVAl PaleoZzoicy | =e r= ire Fig. 4. Measured section of strata exposed in the Ayaguz River basin showing the stratigraphic position of the Ayaguz fossil mammal locality (after Tolochko & Aubekerova 1971). that elsewhere in the Balkash-Alakol basin produce middle Eocene (Irdinmanhan) mammals (on the Shinzhaly River to the south: Didenko-Kislitsina 1990; Lucas et al. 1997). The fossiliferous bed at the Ayaguz River locality is composed of rusty green sandy clays and sandy gravel lenses at the base of a section referred to as ‘“‘Aral svita”’ by Tolochko & Aubekerova (1971). In ad- dition to Franconictis, the following taxa have been reported from this site: the lag- omorphs Sinolagomys cf. S. major Bohlin PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and Desmatolagus cf. D. robustus Matthew & Granger; the castorids Asiacastor ante- cedens Lychev and Steneofiber depereti Mayet; and an indeterminate tragulid artio- dactyl (Musakulova 1971, Erbayeva 1982, Lychev 1982). Desmatolagus ranges in age from Ergi- lian to Shandgolian in China-Mongolia, and is a characteristic Oligocene genus (Russell & Zhai 1987, Qiu & Qiu 1995). The och- otonid Sinolagomys is a characteristic late Oligocene taxon in China and Mongolia, though it has its youngest occurrence (S. pachygnathus) in the early Shanwangian (early Miocene) of China (Qiu & Qiu 1995). Therefore, Erbayeva (1982) assigned the Ayaguz mammal locality a late Oligo- cene age. Lychev (1982, 1987, in Lychev & Au- bekerova 1971) described Asiacastor from localities in eastern Kazakstan that he as- signed a Miocene age, although there is no clear basis for the age assignment. Indeed, Asiacastor is known from the late Oligo- cene Aral local fauna, north of the Aral Sea in western Kazakstan (Lucas et al. 1998). In Asia, the genus Steneofiber is of Oligo- cene and Miocene age (Russell & Zhai 1987). Tragulids range through the Oligo- Miocene boundary, so the indeterminate tragulid from the Ayaguz locality is of no precise biochronological significance. We thus support Erbayeva’s (1982) assignment of a late Oligocene age to the Ayaguz mam- mal locality. Franconictis is known in European early Miocene (MN 1—4) mammal assemblages (Mein 1989, Bruijn et al. 1992, Wolsan 1993, Morlo 1996, Steininger et al. 1996). Its occurrence in eastern Kazakstan thus is an extension of its temporal range back to the late Oligocene. Wolsan’s (in litt. 1997) interpretation of the Kazak specimen of Franconictis as more primitive than Euro- pean Franconictis vireti is consistent with an older age for the Kazak occurrence. Fur- thermore, given that the distribution of Franconictis in Asia is largely unknown, an older age for the genus in Asia than in Eu- VOLUME 111, NUMBER 3 rope is quite plausible. On face value it might suggest an Asian origin of the genus during the Oligocene, and subsequent im- migration to Europe in the early Miocene. Acknowledgments The National Geographic Society (Grant 5412-95) and the Smithsonian Institution’s Charles D. Walcott Fund supported this re- search. R. Tedford, X. Wang, and M. Wol- san provided helpful comments on carni- vore taxonomy and the identification of the specimen reported here. The manuscript was improved by the careful critical re- views of J. Flynn and M. Wolsan. Literature Cited Bowdich, T. E. 1821. An analysis of the natural clas- sification of Mammalia for the use of students and travellers. J. Smith, Paris, 115 pp. Bruijn, H. De, R. Daams, G. Daxner-H6ck, V. Fahl- busch, L. Ginsburg, P. Mein, J. Morales, E. Heinzemann, D. E Mayhew, A. J. Van Der Muelen, N. Schmidt-Kittler, & M. Telles An- tunes. 1992. Report of the RCMNS working group on fossil mammals, Reisensburg 1990.— Newsletters on Stratigraphy 26:65-118. Dashzeveg, D. 1996. Some carnivorous mammals from the Paleogene of the eastern Gobi Desert, Mongolia, and the application of Oligocene car- nivores to stratigraphic correlation.—American Museum Novitates 3179:1—14. . Dehm, R. 1950. Die Raubtiere aus dem Mittel-Miocan (Burdigalium) von Wintershof-West bei Eichs- tatt in Bayern.mAbhandlungen der Bayerischen Akademie der Wissenschaften Mathematisch- naturwissenschaftliche Klasse, Neue Folge 58: 1-141. Didenko-Kislitsina, L. K. 1990. Geologicheskoye raz- vitiye yugo-vostochnovo Kazakhstana v Kai- nozoye [Geological development of southeast- em Kazakstan during the Cenozoic]. Akademi- ya Nauk Kazakhskoy SSR, Institut Gidrogeo- logii 1 Gidrofiziki, Almaty, 49 pp. Emry, R. J., S. G. Lucas, & B. U. Bayshashov. 1997. Brontothere bone bed in the Eocene of eastern Kazakstan.—Journal of Vertebrate Paleontology 17 (supplement to no. 3):44A. Erbayeva, M. A. 1982. Kainozoiskie zaitseobraznye Kazakstana [Cenozoic lagomorphs of Kazak- stan].—Materialy po Istorii Fauny i Flory Ka- zakhstana 8:25-—38. Fischer, G. 1817. Adversaria zoologica—Mémoires de 509 la Société Impériale des Naturalistes de Moscou 5:357-—472. Gabuniya, L. K. 1964. Benarskaya fauna oligotseno- vykh pozvonochnykh [Benara fauna of Oligo- cene vertebrates]. Izdatelstvo ‘‘Metsniereba,”’ Tbilisi, 266 pp. Lucas, S. G., & B. U. Bayshashov. 1996. The giant rhinoceros Paraceratherium from the late Oli- gocene at Aktau Mountain, southeastern Ka- zakhstan, and its biochronological signifi- cance.—Neues Jahrbuch fiir Geologie und Pa- laontologie, Monatshefte 1996:539—548. , R. J. Emry, & B. U. Bayshashov. 1997. Eo- cene Perissodactyla from the Shinzhaly River, eastern Kazakhstan.—Journal of Vertebrate Pa- leontology 17:235—246. , E. G. Kordikova, & R. J. Emry. 1998. Oli- gocene stratigraphy, sequence stratigraphy and mammalian biochronology north of the Aral Sea, western Kazakhstan.—Bulletin of the Car- negie Museum of Natural History 34:313-348. Lychev, G. E 1982. Novyye nakhodki bobrovikh v Pavlodarskoi i Semipalatinskoi oblastyakh [New finds of beavers in the Pavlodar and Sem- ipalatinsk Districts].—Materialy po Istorii Fau- ny i Flory Kazakhstana 8:39—49. . 1987. Novyye svedeniya 0 vymershikh bob- rovykh Prizaysanya i sopredelnovo Sintsizyan- ya [New data on extinct beavers from Prizaysan and adjacent Xinjiang].—Materialy po Istorii Fauny i Flory Kazakhstana 9:69-81. , & P. A. Aubekerova. 1971. Iskopayemyye bobry Kazakhstana [Fossil beavers of Kazakh- stan].—Materialy po Istorii Fauny 1 Flory Ka- zakhstana 5:12—33. Matthew, W. D., & W. Granger. 1924. New Carnivora from the Tertiary of Mongolia.—American Mu- seum Novitates 104:1—9. Mein, P. 1989. Updating of MN zones. Pp. 73—90 in E. H. Lindsay, V. Fahlbusch & P. Mein, eds., European Neogene mammal chronology. Ple- num Press, New York. Mellett, J. S. 1968. The Oligocene Hsanda Gol For- mation, Mongolia: A revised faunal list— American Museum Novitates 2318:1—16. Morlo, M. 1996. Carnivoren aus dem Unter-Miozan des Mainzer Beckens (2. Mustelida, Pinnipedia, Feliformia, Palaeogale).—Senckenbergiana Le- thaea 76:193—249. Musakulova, L. T. 1971. Myestonakhozhdeniye isko- payemykh tragulid v Kazakhstanye [Localities of fossil tragulids in Kazakhstan].—Materialy po Istorii Fauny i Flory Kazakhstana 5:52—56. Qiu, Z., & Z. Qiu. 1995. Chronological sequence and subdivision of Chinese Neogene mammalian faunas.—Palaeoecology, Palaeogeography, Pa- laeoclimatology 116:41—70. Russell, D. E., & R. Zhai. 1987. The Paleogene of 510 Asia: Mammals and _ stratigraphy—Mémoires du Muséum National d’Histoire Naturelle, Sci- ences de la Terre 52:1—488. Steininger, EF FE, W. A. Berggren, D. V. Kent, R. L. Bernor, S. Sen, & J. Agusti. 1996. Circum-Med- iterranean Neogene (Miocene and Pliocene) marine-continental chronologic correlations of European mammal units. Pp. 7—46 in R. L. Ber- nor, V. Fahlbusch, & H-W. Mittmann, eds., The evolution of western Eurasian Neogene mam- mal faunas. Columbia University Press, New York. Tleuberdina, P A., L. T. Abdrakhmanova, & B. U. Bayshashov. 1993. Rannemiotsyenovaya fauna mlyekopitayushikh Dzhungarskovo Alatau (G. Aktau) [Early Miocene mammalian fauna of the Dzhungarian Alatau (Aktau Mountain)].—Ma- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON terialy po Istorii Fauny 1 Flory Kazakhstana 12: 92-115. Tolochko, V. V., & P A. Aubekerova, P. A. 1971. Eot- sen-Pliotsenovyye otlozheniya sredney chasti basseyna r. Ayaguz [Eocene-Pliocene outcrops of the middle part of the Ayaguz River ba- sin].—Geologiya i Poleznye Iskopayemyye Yuzhnovo Kazakstana, 4:154—160. Werdelin, L. 1996. Carnivores, exclusive of Hyaeni- dae, from the later Miocene of Europe and western Asia. Pp. 271—289 in R. L. Bernor, V. Fahlbusch & H-W. Mittmann, eds., The evolu- tion of western Eurasian Neogene mammal fau- nas. Columbia University Press, New York. Wolsan, M. 1993. Phylogeny and classification of early European Mustelida (Mammalia: Carnivora).— Acta Theriologica 38:345—384. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):511—520. 1998. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 6. An intergeneric hybrid, Aglaiocercus kingi X Metallura tyrianthina, from Venezuela Gary R. Graves Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—An intergeneric hybrid hummingbird, Aglaiocercus kingi X Me- tallura tyrianthina, is described. External measurements of the hybrid are in- termediate of those of the parental species. Back plumage iridescence is bluer (511 nm) in the hybrid than in either of the parental species (553-571 nm). This color shift is thought to be caused by a developmental aberrancy or mu- tation which affects melanin granules that produce iridescence in feather ker- atins. Under certain circumstances, interspecif- ic hybridization may be an important source of genetic exchange among avian lineages that may create favorable conditions for rapid and significant evolutionary change (Grant & Grant 1992). From an analysis of data in Panov’s (1989) catalog of avian hy- brids, Grant & Grant (1992) reported that 19.1% (61 of 319) of hummingbird species has hybridized in nature. A surprising 69.2% (36 of 52) of the hybridizing pairs is intergeneric (taxonomy of Sibley & Mon- roe 1990), a finding consistent with Prager & Wilson’s (1975) thesis that interspecific hybridization potential is slowly lost during avian evolution. The true extent of hybrid- ization among hummingbirds, however, is imperfectly known. Panov’s (1989) compi- lation includes numerous poorly document- ed or erroneous records, as did its antece- dent (Gray 1958). Moreover, many new hy- brid combinations have been reported re- cently (e.g., Graves 1990, 1996a, 1998a; Graves & Zusi 1990; Hinkelmann 1996; Weller & Schuchmann 1997). A definitive analysis of hybridization and phyletic retic- ulation must await a robust phylogeny and a systematic survey of purported hybrids, type specimens, and museum collections. Here I describe an intergeneric hybrid com- bination, Aglaiocercus kingi X Metallura tyrianthina. Materials and Methods The unsexed specimen (American Mu- seum of Natural History [AMNH] 146645) was collected by S. Gabaldon in Estado Mérida, Venezuela. The exact locality, ele- vation, and date of collection are unknown. The specimen appears to be a male in sub- definitive plumage as evidenced by the faint striations on the maxillary ramphotheca (see Ortiz-Crespo 1972) and by its elongat- ed tail (Fig. 1 & 2). Five different identifi- cations have been written in ink and pencil (in quotations below) on the two attached AMNH labels since the specimen was cat- aloged in 1927 (in probable order of occur- rence): (a) ““Cyanolesbia’’ [=Aglaiocer- cus]; (b) ‘‘Aglaiocercus ?caudata”’ [=Aglaiocercus kingi caudatus]; (c) ““Me- tallura purpureicauda”’ [=Chalcostigma purpureicauda]; (d) ““?Hybrid?, Aglaiocer- cus caudata X Ramphomicron”’ [=Aglaio- cercus kingi caudatus X Ramphomicron microrhynchum]; and (e) ““Aglaiocercus emmae_ caudata, (melanistic aberration), 512 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Ventral views of male Aglaiocercus kingi caudatus (top), Metallura tyrianthina oreopola (bottom), and a probable hybrid, A. kingi caudatus X M. tyrianthina oreopola (AMNH 146645). Fig. 2. Probable hybrid, Aglaiocercus kingi caudatus X Metallura tyrianthina oreopola (AMNH 146645). VOLUME 111, NUMBER 3 fide J. Berlioz, Apr. 1975” [=Aglaiocercus kingi caudatus}. I compared the oft-identified specimen with series of all species in the subfamily Trochilinae, the typical hummingbirds (Zusi & Bentz 1982, Sibley & Monroe 1990, Bleiweiss et al. 1997), in the collections of the American Museum of Natural History and the National Museum of Natural His- tory (USNM), Smithsonian Institution. For the purpose of hybrid diagnosis, I consid- ered all hummingbirds (Trochilinae; taxon- omy of Sibley & Monroe 1990) that occur in Estado Mérida as potential parental spe- cies (Phelps & Phelps 1958, Meyer de Schauensee & Phelps 1978) (Appendix 1). In addition, I compared the specimens di- rectly with the holotypes of Chalcostigma purpureicauda (AMNH 483931), Lesbia ortoni (AMNH 156651), Zodalia thaumas- ta (USNM 173911), and Aeronympha pro- santis (Field Museum of Natural History, FMNH 11852), and with notes, photo- graphs, and videotape of the holotype of Heliangelus zusii (Academy of Natural Sci- ences of Philadelphia, ANSP 159261). Color descriptions given in Appendix 2 were made under natural light. I evaluated the color of dorsal plumage (center of back) and the ventral surfaces of the rectrices with a reflectance spectrophotometer equipped with a 11.0 mm aperture (Color Mate Col- orimeter, Milton Roy). The colorimetric characters were described in terms of op- ponent-color coordinates (L, a, b) (Hunter & Harold 1987). This system is based on the hypothesis that signals from the cone receptors in the human eye are coded by the brain as light-dark (L), red-green (a), and yellow-blue (b). The rationale is that a color cannot be red and green or yellow and blue at the same time. Therefore “‘redness’’ and ““greenness”’ can be expressed as a single value a, which is positive if the color is red and negative if the color is green. Likewise, “‘yellowness”’ or “‘blueness”’ is expressed by b for yellows and —b for blues. The third coordinate L, ranging from 0 to 100, describes the “‘lightness”’ of color; low val- 513 ues are dark, high values are light. In other words, the more light reflected from the plumage the higher the L value will be. It should be noted that visual systems in hum- mingbirds (e.g., Goldsmith & Goldsmith 1979) differ significantly from those of hu- mans. The relevance of opponent color co- ordinates to colors perceived by humming- birds is unknown. Dominant wavelengths reflected from plumage surfaces are listed for comparison. Data in Table 1 were com- piled from the averages of five independent measurements (specimen moved from ap- erture between trials) for each plumage area per specimen. Measurements of wing chord, bill length (from anterior extension of feathers), and rectrix length (from point of insertion of the central rectrices to the tip of each rectrix) were taken with digital calipers and round- ed to the nearest 0.1 mm (Table 2). Mea- surements and least squares regression lines were projected on bivariate plots to illus- trate size differences (Wilkinson 1989). There are three alternatives to consider— the specimen represents an aberrant color morph of A. kingi or some other species, a hybrid, or an undescribed species. The specimen differs significantly in size and shape from all species in Appendix 1. In particular, the rectrices of the specimen are considerably wider, flatter in cross section, and more iridescent on the ventral surfaces than in A. kingi, indicating that it is not sim- ply a melanistic example of that species as suggested by Berlioz on the specimen label. Because hybrids have no standing in zoo- logical nomenclature, the burden of proof rests on the investigator to refute this pos- sibility before bestowing species status on a unique specimen. Because the evidence points to hybridization, I refer to the spec- imen as a hybrid in the remainder of the paper. The diagnosis was approached in a hi- erarchical manner. The presumed parental species of the hybrid first were hypothe- sized through the comparative analysis of plumage pattern, as well as from feather 514 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Ranges and means (+standard deviation) of opponent color coordinates (L, a, b) and dominant wavelength reflected from dorsal plumage (center of back) and the ventral surface of rectrices in male Aglaio- cercus kingi caudatus, Metallura tyrianthina oreopola, and their probable hybrid (AMNH 146645). A. kingi M. tyrianthina Variables (n = 12) (n = 12) Hybrid Back plumage L (Lightness) (L) 23.6—30.1 21.5—26.6 24.1 PH 2e Wess DAD r= eA) a (Red [+]/Green [—]) (a) —14.8-(—6.6) —7.1-(—0.6) —7.9 =10:9 = 2-9 S37 3E i183 b (Yellow [+]/Blue [—]) (b) 15.4—20.2 12.6—20.9 2.9 17.8 + 1.4 17.3 2 DB Dominant Wavelength (nm) 553.1-563.8 562.8—570.5 511.3 Sas) 2 SD) 566.7 + 2.6 Ventral surface of rectrices L (Lightness) (L) 18.8—21.1 20.3—23.9 20.6 2 On OM Die = NAD) a (Red [+]/Green [—]) (a) 3.8-5.9 11.5-17.0 10.6 49 + 0.6 14.6 + 1.6 b (Yellow [+]/Blue [—]) (b) —4.3-1.5 5.5-14.9 =7/.\l —0.9 + 1.9 OS) a6 DY) Dominant Wavelength (nm) 487.2-652.1 588.9-622.7 541.0 SAO 22 47/7) 602.5 + 11.2 and bill shape. The restrictive hypothesis then was tested with a quantitative analysis of size and external proportions. Concor- dance of results is regarded as strong sup- Table 2.—Ranges and means (+standard deviation) of measurements (mm) of males of Aglaiocercus kingi caudatus (subdefinitive plumage, see Appendix 2), Metallura tyrianthina oreopola, and their presumed hybrid (AMNH 146645). port for the hypothesis (Graves 1990, Graves & Zusi 1990). Results and Discussion Plumage characters.—Salient characters of the hybrid that permit its parental species to be identified include: (a) moderately elongated outer rectrices (fork depth = 23.7 mm), nearly flat in cross section; (b) un- marked rectrices exhibiting metallic irides- A. kingi M. tyrianthina cence on the dorsal and ventral surfaces; (c) Character (n = 21) (n = 20) Hybrid Sig i short tibial plumes (not extending to hal- Wing chord 59.7-63.6 54.0-61.5 61.4 yx); and (d) short straight bill (11.1 mm). GiEsi== sO Sell 25 DP ies : Two species in the pool of potential paren- Bill 11.0-13.6 91-106 11-1 1 d A = OF Ao aati tal species (Appendix 1) possess elongate i i > : - Rectrix | DD BEDS 33.7389 "32, ‘alls Cengiheiitect > ae 241 +08 261 £16 tus underwoodii and Aglaiocercus kingi. Rectrix 2 WO.O-33.6 34.3-41.3 36.9 Ocreatus can be deleted from the list of Sills} se 37.9 + 1.8 possibilities because the hybrid lacks evi- Rectrix 3 38.9-46.7 36.0-43.6 43.1 dence of spatulate rectrices or lengthened 42.8 + 1.9 Sie) 25 1 tibial plumes. Aglaiocercus kingi is thus Rectrix 4 53.7-61.5 38.4-45.3 49.0 identified as one of the parental species. 20) Zot AUS = 2:1 Determination of the other parental spe- Ege 3 G75 L027 37.9-46.3 55.8 cies is equally straightforward. The inten- 3 oe) ADD, se 7) sity of the metallic iridescence reflected VOLUME 111, NUMBER 3 600 S4) @ oOo 560 540 520 Dominant Wavelength (nm) of Back Plumage 500 Lightness (L) of Back Plumage Fig. 3. 515 Yellowness (b) of Back Plumage O -10 -5 O 5 10 15 Yellowness (b) or Blueness (-b) of Ventral Surface of Rectrices Bivariate plots of spectrophotometric data from male hummingbirds: Aglaiocercus kingi caudatus (circles); Metallura tyrianthina oreopola (diamonds); and a probable hybrid, A. kingi caudatus * M. tyrianthina oreopola (triangle; AMNH 146645). from the ventral surfaces of the hybrid’s rectrices is matched or exceeded only in Metallura tyrianthina. Details of plumage pattern and feather shape are sufficient to suggest that the parentage of the hybrid is Aglaiocercus kingi X Metallura tyrianthina (see Appendix 2). None of the other species in Appendix 1, considered two at a time, can account for the characters observed in the hybrid. In particular, the ventral rectri- cial surfaces of the hybrid are metallic red- dish-purple as opposed to dull black or pur- plish-black in both Ramphomicron micro- rhynchum and A. kingi, effectively elimi- nating this pair of species from contention. The question of plumage color.—Irides- cence in hummingbirds is caused by the in- _terference of light reflected from the upper and lower surfaces of gas-filled vacuoles in melanin granules in the keratin of feather barbules, which are compactly stacked in 7—15 layers in the barbule keratins (Dorst 1951; Greenewalt et al. 1960a, 1960b; Lu- cas & Stettenheim 1972). Carotenoid pig- ments have not been extracted from irides- cent feathers. Employing transmission electron microscopy and micro-spectropho- tometry, Greenewalt et al. (1960a, 1960b) found melanin granules to be elliptical in shape, about 2.54 long, 1.54 wide, and 0.15y thick. Briefly summarized, they found that granules contain a fairly uniform layer of gas-filled vacuoles that resemble a monolayered foam. The melanin matrix and gas-filled vacuoles have refractive indices of ~2.0 and 1.0, respectively. The color of iridescence varies according to the thick- ness of the granule and the amount of gas in the vacuoles. Iridiscent colors change from blue to green to orange and finally to red, as the effective refractive index of granules advances from 1.45 to 1.90 (figure 4 of Greenewalt et al. 1960a). Melanin granules in noniridescent parts of feathers lack vacuoles. The pattern of bluish-green iridescence in the hybrid corresponds precisely to that of green iridescence in the parental species, suggesting a single mutational or develop- mental aberrancy that affects plumage col- or. The dominant wavelength reflected from dorsal plumage is shorter in the hybrid (511 nm) than in the parental species: Aglaio- cercus kingi (553-564 nm) and Metallura tyrianthina (563-571 nm) (Table 1, Fig. 3). The premise that “‘hybridization produces no traits characteristic of genera or species other than those involved in the particular cross”’ (Banks & Johnson 1961:3) was ex- tended to spectrophotometric measures of 516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 65 60 Wing Length 55 50 20 40 60 80 Length Rectrix 5 40 35 30 Length Rectrix 1 25 20 20 30 40 Length Rectrix 2 Length Rectrix 1 30 40 50 Length Rectrix 4 Fig. 4. Bivariate plots of mensural characters of male hummingbirds: Aglaiocercus kingi caudatus (triangles); Metallura tyrianthina oreopola (circles); and a probable hybrid, A. kingi caudatus X M. tyrianthina oreopola 100 60 120 70 14 13 12 Bill Length 11 10 20 40 60 80 100 120 Length Rectrix 5 40 w@ (o) () Oo Length Rectrix 1 i) a 20 35 40 45 50 Length Rectrix 3 40 ie) on ie) Oo Length Rectrix 1 ie) 6) a Length Rectrix 5 (diamond; AMNH 146645). Least squares regression lines are projected for comparison. er VOLUME 111, NUMBER 3 plumage color (Graves 1996b). Both as- sumptions are violated in the present ex- ample. Elucidation of the micro-structure and spectrophotometric properties of melanin granules in Aglaiocercus kingi, Metallura tyrianthina, and the hybrid is beyond the scope of this paper. Several observations on hummingbird plumages, however, are worth noting. A variety of plumage aber- rations, unassociated with hybridization, has been observed in hummingbirds, in- cluding leucism, albinism, schizochroism, erythrism, and melanism (Salvin 1892, Banks & Medina 1963, Greenway 1978, Graves 1998b). Subtle within-population variation in iridescent color is commonly observed whenever large series of species are assembled. Although post-mortem ef- fects may be partially responsible in some cases (Graves 1986, 1991), most of the ob- served variation in iridescence among in- dividuals, factoring out the effects of sex and age (see Bleiweiss 1992), is due to ge- netic and developmental factors. Pro- nounced color shifts of the magnitude ob- served in this hybrid are rare but not un- known (e.g., Salvin 1892, Greenway 1978). The example described here seems to be the first in which a hybrid hummingbird exhib- its a major plumage aberrancy. External measurements.—One of the guiding principles of hybrid diagnosis is that hybrids are not larger or smaller than their parental species (Graves 1990). Mor- phological luxuriance or dwarfism in hybrid hummingbirds has not been recorded. Male Aglaiocercus kingi and Metallura tyrianthi- na are similar in bill length (cumulative range, 9.1—13.6 mm) and wing chord (cu- mulative range, 54.0—63.6 mm), but differ markedly in tail size and shape (Table 2, Fig. 4). Bivariate plots of rectrix length of the parental species exhibit positive (1 vs. 2) or negative (1 vs. 3, 1 vs. 4, 1 vs. 5) allometry. Except for rectrix 3, measure- ments of the hybrid fall between the char- acter means for A. kingi and M. tyrianthina, and, in several cases, approximate the val- S17) ues predicted by least squares regression on bivariate plots (Table 2, Fig. 4). In sum- mary, plumage pattern, distribution and in- tensity of iridescence, rectrix shape, and mensural characters provide strong support for the hypothesis of hybridity (Aglaiocer- cus kingi X Metallura tyrianthina). Previous records.—A hybrid of Aglaio- cercus kingi and Metallura tyrianthina was reported once before by Meyer de Schauen- see (1947:108), who described a specimen (No. 134) obtained in Bogota, circa 1909, from the Brother Nicéforo Maria collection: “... fore-crown glittering brassy green, hind crown and back dark bluish green, rump and upper tail coverts bluer; chin dusky, throat patch shaped as in Metallura tyrianthina but blue instead of green; breast dark bluish green, bases and edges of the feathers buffy; belly dark green, the bases of the feathers white, showing through and giving a some- what barred appearance; tail purple, deeply forked, the outermost tail feathers 50 mm., the central ones 30 mm., wing 61 mm., culmen 12.5 mm.” The brief description of Nicéforo’s spec- imen differs in minor details from the Ven- ezuelan specimen (AMNH 146645). The two specimens are similar in size. Nicéfo- ro’s specimen possesses a bluish gorget as might be expected in an adult male hybrid of Metallura t. tyrianthina and Aglaiocer- cus k. kingi from the Cordillera Oriental of the Colombian Andes. Whereas I charac- terized the back color of the Venezuelan specimen as “greenish-blue,’’ Meyer de Schauensee used the term “bluish-green’”’ for Nicéforo’s specimen. This and other dis- crepancies might reflect semantics or real differences in color. Unfortunately, the whereabouts of Nicéforo’s specimen is un- known, although another mentioned in Meyer de Schauensee’s paper was deposit- ed in the Academy of Natural Sciences of Philadelphia (Nicéforo no. 148, now ANSP 159261; Graves 1993). Acknowledgments I thank Richard Banks, Robert Bleiweiss, Kenneth C. Parkes, and Richard Zusi for critiques of the manuscript. I thank the cu- 518 rators and staffs of the American Museum of Natural History, New York, the Field Museum of Natural History, Chicago, and the Academy of Natural Sciences of Phil- adelphia, for permitting me to examine specimens in their care and for specimen loans. Photographic prints were provided by Smithsonian photographic services. Mu- seum work was supported by the Alexander Wetmore Fund and the Department of Ver- tebrate Zoology, National Museum of Nat- ural History, Smithsonian Institution. Literature Cited Banks, R. C., & N. K. Johnson. i961. A review of North American hybrid hummingbirds.—Con- dor 63:3—28. , & D. R. Medina. 1963. An albinistic Anna Hummingbird.—Condor 65:69—70. Bleiweiss, R. 1992. Reversed plumage ontogeny in a female hummingbird: implications for the evo- lution of iridescent colours and sexual dichro- matism.—Biological Journal of the Linnean So- ciety 47:183-195. , J. A. W. Kirsch, & J. C. Matheus. 1997. DNA hybridization evidence for the principal lineages of hummingbirds (Aves: Trochilidae).—Molec- ular Biology and Evolution 14:325—343. Dorst, J. 1951. Recherches sur la structure des plumes des Trochilidés——Meémoires du Muséum Na- tional D’Histoire Naturelle (Série A. Zoologie) 1:125—260. Goldsmith, T. H., & K. M. Goldsmith. 1979. Discrim- ination of colors by the black-chinned hum- mingbird, Archilochus alexandri.—Journal of Comparative Physiology A 130:209—220. Grant, P. R., & B. R. Grant. 1992. Hybridization of bird species.—Science 256:193—-197. Graves, G. R. 1986. Systematics of the Gorgeted Woodstars (Trochilidae: Acestrura).—Proceed- ings of the Biological Society of Washington 99:218—-224. . 1990. Systematics of the ““green-throated sun- angels” (Aves: Trochilidae): valid taxa or hy- brids?—Proceedings of the Biological Society of Washington 103:6—25. . 1991. Taxonomic status of the Sword-billed Hummingbird (Ensifera ensifera caerules- cens).—Bulletin of the British Ornithologists’ Club 111:139-140. . 1993. Relic of a lost world: a new species of sunangel (Trochilidae: Heliangelus) from Bo- gota.—Auk 110:1-8. . 1996a. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 2. Hybrid origin of Erioc- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nemis soderstromi Butler—Proceedings of the Biological Society of Washington 109:764—769. 1996b. Hybrid wood warblers, Dendroica striata X Dendroica castanea (Aves: Fringilli- dae: Tribe Parulini) and the diagnostic predict- ability of avian hybrid phenotypes.—Proceed- ings of the Biological Society of Washington 109:373-—390. , 1998a. Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 5. Probable hybrid origin of Amazilia distans Wetmore & Phelps.—Pro- ceedings of the Biological Society of Washing- ton 111:28—34. , 1998b. Taxonomic notes on hummingbirds (Aves: Trochilidae). 1. Eriocnemis dyselius El- liot, 1872 is a melanistic specimen of Eriocnem- is cupreoventris (Fraser, 1840).—Proceedings of the Biological Society of Washington 111: 420-424. , & R. L. Zusi. 1990. An intergeneric hybrid hummingbird (Heliodoxa leadbeateri X Helian- gelus amethysticollis) from northern Colom- bia.—Condor 92:754—760. Gray, A. P. 1958. Bird hybrids. Commonwealth Agri- cultural Bureaux, Bucks, England, 390 pp. Greenewalt, C. H., W. Brandt, & D. D. Friel. 1960a. Iridescent colors of hummingbird feathers.— Journal of the American Optical Society 50: 1005-1016. . 1960b. The iridescent colors of hummingbird feathers.—Proceedings of the American Philo- sophical Society 104:249—253. Greenway, J. C., Jr. 1978. Type specimens of birds in the American Museum of Natural History. Part 2.—Bulletin of the American Museum of Nat- ural History 161:1—305. Hinkelmann, C. 1996. Evidence for natural hybridi- sation in hermit hummingbirds (Phaethornis spp.).—Bulletin of the British Ornithologists’ Club 116:5-14. Hunter, R. S., & R. W. Harold. 1987. The measurement of appearance. 2nd edition. Wiley, New York, 411 pp. Lucas, A. M., & P. R. Stettenheim. 1972. Avian anat- omy. Integument, Part 2.—United States De- partment of Agriculture, Washington, D.C., Ag- ricultural Handbook 362:341—750. Meyer de Schauensee, R. 1947. New or little-known Colombian birds.—Proceedings of the Acade- my of Natural Sciences of Philadelphia 99:107— 126. , & W. H. Phelps, Jr. 1978. A guide to the birds of Venezuela. Princeton University Press, 424 Pp. Ortiz-Crespo, E I. 1972. A new method to separate immature and adult hummingbirds.—Auk 89: 851-857. Panov, E. N. 1989. Natural hybridisation and etholog- VOLUME 111, NUMBER 3 ical isolation in birds (in Russian). Nauka, Mos- cow, 510 pp. Phelps, W. H., & W. H. Phelps, Jr. 1958. Lista de las aves de Venezuela con su distribuci6n. Tomo 2, Parte 1. Editorial Sucre, Caracas, 317 pp. Prager, E. M., & A. C. Wilson. 1975. Slow evolution- ary loss of the potential for interspecific hybrid- ization in birds: a manifestation of slow regu- latory evolution.—Proceedings of the National Academy of Science USA 72:200—204. Salvin, O. 1892. Catalogue of the birds in the British Museum, Vol. 16, London, 703 pp. Sibley, C. G. & B. L. Monroe, Jr. 1990. Distribution and taxonomy of birds of the world. Yale Uni- versity Press, New Haven, Connecticut, 1111 Pp. Weller, A.-A., & K.-L. Schuchmann. 1997. The hybrid origin of a Venezuelan Trochilid, Amazilia dis- tans Wetmore & Phelps 1956.—Ornithologia Neotropical 8:107—112. Wilkinson, L. 1989. SYSTAT: the system for statistics. SYSTAT, Inc., Evanston, Illinois, 822 pp. Zimmer, J. T. 1952. Studies of Peruvian birds. No. 62. The hummingbird genera Patagona, Sappho, Polyonymus, Ramphomicron, Metallura, Chal- costigma, Taphrolesbia, and Aglaiocercus.— American Museum Novitates 1595:1—29. Zusi, R. L., & G. D. Bentz. 1982. Variation of a muscle in hummingbirds and swifts and its systematic implications.—Proceedings of the Biological Society of Washington 95:412—420. Appendix 1 Species of hummingbirds that occur in Estado Mé- rida, Venezuela: Campylopterus falcatus, Colibri thal- assinus, C. coruscans, Klais guimeti, Lophornis delat- trei, L. stictolophus, Chlorestes notatus, Chlorostilbon mellisugus, C. poortmani, Thalurania furcata, Hylo- charis cyanus, Chrysuronia oenone, Amazilia versi- color, A. fimbriata, A. viridigaster, Chalybura buffonii, Heliodoxa leadbeateri, Sternoclyta cyanopectus, Coe- ligena coeligena, Ocreatus underwoodii, Aglaiocercus kingi, Heliomaster longirostris, Chaetocercus jourda- nil. Appendix 2 Comparative description of plumages of male Aglaiocercus kingi caudatus, Metallura tyrianthina or- eopola, and their presumed hybrid, AMNH 146645. The molts and plumages of male Aglaiocercus spp. are incompletely known. Young males (>6 months?) ac- quire a plumage that differs from the definitive plum- age of adult males. This subdefinitive plumage is char- acterized by shorter outer rectrices, an incompletely developed crown patch (8 of 20 examined), and rem- nants of a white rump patch (see Zimmer 1952). One quarter (5 of 20) of the males in subdefinitive plumage 519 retain a few striations on the maxillary ramphothecum, a character usually interpreted as a sign of immaturity (Ortiz-Crespo 1972). The descriptions of Aglaiocercus kingi given below refer to the subdefinitive plumage. Descriptions of structural colors are unusually sub- jective, as color seen by the observer varies according to the angle of inspection and direction of light. For this reason I use general color descriptions. The dark bluish-green crown of young kingi is re- placed (from anterior to posterior) by an ovate crown patch composed of brilliant bluish-green feathers. The hindneck, back, and rump are dark green; feathers are gray, tipped with green. Upper-tail coverts are bluish- green. A few white feathers form an indistinct patch on the lower back. The dorsal plumage of tyrianthina is dark dusky green, brighter on the crown, and with coppery high- lights on the lower back and rump. Feathers are gray, banded subterminally with coppery-green, and tipped broadly with dark green. When viewed head-on in di- rect light, plumage posterior to the midcrown region appears sooty black. Immature tyrianthina lack a con- trasting rump patch. Under a diffuse light source, the dorsum of the hy- brid is a rich greenish-blue (paler on the crown), a color that is distinctly different from that of the pre- sumed parental species. Feathers on the left side of the forecrown are discolored, possibly by a preservative chemical. Dorsal feathers are dark gray, tipped with greenish-blue. Crown feathers are not modified as in adult kingi. When light is reflected obliquely (>90 from the observer), the dorsal plumage appears purple; when viewed head-on the hindcrown, back, and rump appear black. A few rump feathers are tipped with buff. The ventral plumage of kingi is medium green ex- hibiting subdued iridescence. A few small shining green disks occur on the throat of more mature indi- viduals. The barbs of ventral feathers are narrowly tipped with buff or grayish-buff, especially along the midline of the abdomen. Some males in juvenile and subdefinitive plumage (e.g., AMNH 484067) have a white or buffy-white line extending from the base of the bill posterior to below the eye. Downy vent feath- ers are dark gray tipped with white or pale gray. Un- dertail coverts are dark green broadly edged with buff. Tibial feathers are short (extending half way to the hallux from the tibiotarsal joint), dark olive-gray and narrowly tipped with scattered grayish-buff barbs. The venter of tyrianthina is dark dull green; feathers are tipped with buff or grayish-brown, especially along the midline. Feather tipping imparts a mottled appear- ance to the underparts. A narrow ovate gorget extends from the chin to the upper breast in tyrianthina. When view head-on in direct light, the auriculars and sides of the throat appear matte black and contrast greatly with the brilliant green gorget. Subadult males have a buffy line extending from the bill to below the eye. 520 Vent feathers are dark gray tipped with white or pale gray. Under-tail coverts are buff with a large broadly lanceolate spot (coppery- or bronzy-red) along the midline. Tibial feathers (dark gray tipped with buff) extend to the base of the hallux. The venter of the hybrid is similar in pattern to kin- gi, but with the green portions replaced with bluish- green (paler than dorsum). A small brilliant feather occurs along the midline at midthroat (deep blue tip separated from the gray base by narrow blending bands of light blue and coppery-gold). A buffy-white stripe extends from the bill to below the eye (Fig. 2). Vent feathers are dark gray tipped with white. Undertail co- verts are buff with a lanceolate subterminal spot (pur- ple) near the midline. Tibial feathers (dark gray broad- ly tipped with buff) of the hybrid extend about halfway to the hallux, but may have been damaged by knotting of the specimen label string. The remiges of kingi are brownish-black faintly tint- ed with purple. The outer vanes of the primary coverts and the innermost secondaries are edged with shining green or bluish-green; secondary coverts are broadly tipped with bluish-green. The remiges of tyrianthina are very similar in color but faintly tinted with bronze or olive. Wing coverts and the innermost secondaries are bronzy-green. The remiges of the hybrid resemble those of kingi. Wing coverts and innermost secondaries are purple tipped with dark bluish-green. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The tail of kingi is deeply forked. The basal portions of the rectrices that are obscured in the folded tail are black. The exposed dorsal sections of the inner rectri- ces (1—4) are deep brilliant purple tipped with bluish- green. The outermost rectrices (5) lack bluish-green tips, are greatly elongated, narrow (S—6 mm wide, 25 mm from tip), and bowed in cross-section. Inner rec- trices are smoothly tapered; rectrix 5 is bluntly tipped. Ventrally, the vanes are dull purplish-black. The ra- chises in kingi are blackish-brown dorsally, medium brown ventrally. The tail of tyrianthina is shallowly forked. Rectrices are wide (10-11 mm), nearly flat in cross-section, abruptly truncate at the tip, and metallic coppery-red above and below. Rachises are dark brown above and below. The tail of the hybrid is moderately forked. Feather size and shape are intermediate between kingi and ty- rianthina. The outermost rectrices (5) are slightly bowed in cross-section (ca. 9.3 mm at widest point). Rectrices are metallic reddish-purple, above and be- low, the innermost (1—2) are diffusely tipped with pur- ple. This iridescence, especially from the ventral sur- faces, is similar in visual essence to that of tyrianthina. Rachises are dark brown above, medium brown ven- trally. Bill color is black in kingi, tyrianthina, and hybrid. In dorsal profile, the bills of both parental species are abruptly tapered, more so in kingi. The bill profile of the hybrid is similar to that of tyrianthina. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):521—530. 1998. A new species of Alsodes (Amphibia: Anura: Leptodactylidae) from southern Chile J. Ramon Formas, César Cuevas, and José Nufiez Instituto de Zoologia, Universidad Austral de Chile, Casilla 567, Valdivia, Chile Abstract.—A new species of frog, Alsodes kaweshkari, is described from the temperate Nothofagus forests of South America in Southern Chile. From the karyological point of view this species is included in the monticola group (2N = 26) of the genus Alsodes. The genus Alsodes Bell 1843, is distrib- uted in central and southern Chile and along the eastern slope of the Andes, in Argentina (south of Mendoza city). Frost (1985) in- cluded the following species in the genus: A. barrioi, A. gargola, A. illotus, A. laevis, A. montanus, A. monticola, A. nodosus, A. pehuenche, A. tumultuosus, A. vanzolinii, and A. verrucosus. Formas (1989) studied the identity and synonymy of the Chilean frog Eupsophus vittatus and concluded that this species must be included in the genus Alsodes as A. vittatus. The taxonomic status of A. illotus is not clear (Cei 1980) and ac- cording to Frost (1985) A. laevis has not been collected since its description by Phi- lippi (1902). The most remarkable charac- teristic of the males of this genus is the presence of thorny structures on the fingers and round spiny patches on the chest. During the course of collecting frogs in southern Chile we found a new species of Alsodes which is here described. Moreover, we describe the karyotype and C-banded chromosomic pattern of the new taxon. The bands of hemoglobin of the new species were characterized electrophoretically and compared with those of A. monticola (type species of the genus). Methods and Materials Specimens were deposited in the Instituto de Zoologia, Universidad Austral de Chile (IZUA). Adults were measured with a dial caliper to the nearest 0.1 mm according to Cei (1962). The following measurements were taken: snout—vent length (SVL), head length, head width, nostril-snout distance, tibia length, and foot length. Internarial dis- tance was measured according to Cei (1980) and eye diameter according to Duellman (1970). The chromosomal characteristics were obtained from the holotype (IZUA 1624) according to the following methods: the an- imal was treated with 0.1% colchicine for two hours, then anaesthetized with diethyl ether, cut open ventrally under sterile con- ditions and its intestine carefully removed. Metaphase plates were obtained by squash- ing intestinal epithelium fragments that were hypotonically treated with distilled water, then fixed in acetic-alcohol (1:3), and finally placed in 45% acetic acid. Small tis- sue fragments were squashed between a glass slide and cover slip and dipped in liq- uid nitrogen; thereafter, the cover slip was removed with a razor blade to allow chro- mosomes to air dry. After 3 days, the chro- mosomes were stained for 15 minutes in Sorensen’s phosphate buffer (pH 6.8), con- taining 4% Giemsa solution (Formas 1991). Centromeric positions were determined ac- cording to Levan et al. (1964). Secondary constrictions were included in the measure- ments. Chromosomes were stained to reveal C-band pattern position (Sumner 1972). Microscopic slides and the specimen (IZUA S22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Alsodes kaweshkari Holotype, IZUA 1624. 1624) were deposited in the amphibian col- lection of the Instituto de Zoologia, Univ- ersidad Austral de Chile. Hemoglobin was obtained according to the following methodology: blood was col- lected by cardiac puncture from the holo- type and a specimen of Alsodes monticola (control species) using a sterile syringe with 3.8% of sodium citrate as anticoagulant; the red cells were washed 3 times with 0.85% NaCl and centrifuged at 700 g for 5 minutes at room temperature (20°C). The packed cells were hypotonically treated with dis- tilled water (10 volumes) for 30 minutes and centrifuged at 3500 g for 10 minutes. The supernatant was stored at —70°C. The hemolyzate was treated according to the method described by Muir (1981) with modifications. We used a minicolumn (5 X 70 mm) of Sephadex G-50 equilibrated with 1.0 mM of potassium phosphate (pH 7.0). Fractions of 500 wl were collected and their absorbances measured at 410 nm, the wave length at which the porphyrin ring of the hemoglobin molecule absorbs specifi- cally. Those fractions with the highest ab- sorbance value were used for the electro- phoretic analysis of the hemoglobin of the M. Nowaste . 97 2cm new species and A. monticola. To proceed with the electrophoresis the samples were denatured by heating (100°C, 5 minutes) in presence of 2% of betamercaptoethanol. Electrophoresis of the hemoglobins from Alsodes kaweshkari and A. monticola was carried out individually in vertical slab gels of polycrilamide (15%) in denaturant con- ditions at 20 mA for 20 hours (Laemmeli 1970). Gels were stained with Coomassie Blue R-250 for 2 hours and destained with acetic acid (7%). The molecular weight of the hemoglobin bands was estimated using a protein standard for molecular weight (Sigma Co). Systematics Alsodes kaweshkari, new species. Fig. 1 Holotype.—IZUA 1624, an adult male collected by J. Ram6n Formas on 8 March 1995, at Puerto Edén (49°8'S, 74°25'W, 10 m), Wellington Island, Ultima Esperanza Province, Magallanes and Chilean Antartic Region (XII Regién), Chile (Fig. 2). Paratype.-—IZUA 1625, an adult male collected by Pablo Corti on 21 Oct 1995, VOLUME 111, NUMBER 3 46° Peninsula de Taitao Sf B (e) oO u O > ¢ Q OCEANO Puerto Isla a Wellington ~—-” Fig. 2. Type locality of Alsodes kaweshkari. at Seno Huemules (48°43’'5S, 74°25’W, 8 m), Ultima Esperanza Province, Magallanes and Chilean Antartic Region (XII Region), Chile (Fig. 2). Diagnosis.—Alsodes kaweshkari can be distinguished from its congeneric species by the following combination of characters: SVL 56.5—62.2 mm; toes totally fringed; webbing of feet present between all toes, but reduced; dorso-lateral surfaces granular; skin around vent and posterior thighs gran- ular. Description.—Based on the type series. 523 ) REPUBLICA ? ARGENTINA Seno Huemules Body robust, arms and legs well developed. Head depressed, slightly wider than long, its length 29% of snout—vent length. Snout truncated from above, rounded in lateral profile; canthus rostralis slightly rounded, loreal region concave in cross section; nos- trils anterolateral, midway between tip of snout and anterior border of eye; eye di- ameter greater than distance between eye and nostril; internarial distance less than in- terorbital distance. Tympanum absent; post- ocular fold evident, reaching insertion of arm. Tongue rounded, without notch at tip. 524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SSP rad ey j iis Fig. 3. Morphological details of the holotype. (A) Palmar view of the right hand. (B) Plantar view of the right foot. Choanae oval-shaped; dentigerous process- es of vomer between choanae. Forelimbs of males robust. Fingers in order of increasing length: II, I, IV, II. Webbing of hand ab- sent. Fingers long with moderately globular tips. Inner palmar tubercle ovoid, outer pal- mar tubercle rounded; subarticular tubercles rounded, (Fig. 3A). First finger with strong thorny excrescences; second finger with narrow band of spines (Fig. 4B). Palmar VOLUME 111, NUMBER 3 S25) 5mm Fig. 4. Secondary sexual characters of the holotype. (A) Ventral view of the chest and throat. (B) Dorsal view of the left hand. surface with few thorns irregularly distrib- uted. Spines also present at lower border of mandible (Fig. 4A). Toes long, thick and fringed; in order of increasing length: I-II- III-V-IV; tips moderately rounded. Webbing present, thick, but reduced. Inner metatarsal tubercle oval, elongate; outer tubercle smaller than the inner, oval. Tarsal fold present; reaching middle of tarsus. Flanks and dorsal surface granular; ventral skin smooth with minute granules; skin around vent and posterior thighs granular. Chest of males with two bilateral rounded black patches of keratinous spines (Fig. 4A). Dor- Table 1.—Measurements (mm) of the type series of Alsodes kaweshkari. Holotype Paratype IZUA 1624 IZUA 1625 Character male male Snout—vent length 62.2 59.9 Head length 17.7 16.4 Head width 24.2 22.8 Eye diameter 7.0 6.7 Nostril—snout distance 4.0 3.6 Internarial distance So 7/ 5.4 Tibia length MA 27.4 Foot length 33.8 30.9 sal surfaces of head and areas below dor- solateral fold with granules. The measurements of the holotype and the paratype are given in Table 1. Coloration.—In alcohol, dorsal surfaces of ground, arms and legs dark gray. Venter whitish, gular area dark gray. Arms and legs whitish ventrally. The granular surface around posterior thighs dark gray. In life, dorsal surface, arms and legs brown. Belly whitish yellow. Areas around nostrils, lips and eyes light brown. The iris with a cop- pery reticulum. Distribution and ecology.—The species is known from two Chilean localities (Puer- to Edén and Seno Huemules) (Fig. 2). The type locality (Puerto Edén) is a small beach of the Wellington Island, near a small fish- ing town. There are small forests (Notho- fagus betuloides, Embothrium coccineum, Maytenus magellanicus, and Drymis win- teri) surrounded by typical tundra with plants of the genera Donatia, Astelia, and Azorella and rushes of the families Cyper- aceae and Juncae. This area is situated in the oceanic cold temperate region (di Castri 1968). The annual mean temperature of this region is 8.8°C; the relative humidity is 526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ey i a ax BA aust 1 eA if 11 12 Mon cs 8 a 8 9 10 Bk aks} 10 um } 5 $ = & a1 . 3 4 5 * A 13 Fig. 5. Chromosomes of the holotype. (A) Karyotype. (B) C-banded karyotype. 87%, and the rain fall ranges from 2000— 2500 mm per annum. The frog was col- lected under a log between the forest and the tundra. The male holotype shows its re- markable thorny excrescences on the chest and fingers (Fig. 4A, B). The following am- phibians were also collected in this area: Bufo variegatus, Batrachyla antartandica and Eupsophus calcaratus. Lynch (1975) pointed out the presence of Atelognathus VOLUME 111, NUMBER 3 Table 2.—Relative length, arm ratio (mean and stan- dard deviation), and type of chromosomes (m = meta- centric; sm = submetacentric; st = subtelocentric) of metaphase chromosomes of Alsodes kaweshkari. Rel- ative length was calculated according to Bogart (1970). Arm ratio was calculated by dividing the short arm into the long arm. *Chromosomes with secondary con- strictions. Pair n° Relative length Arm ratio Type ee 148.72 + 18.32 1.97 + 0.20 sm 135.58 = 18.76 1.71 = 0.63 sm. 3 126.80 + 10.50 4.69 + 0.48 st 4* 116.74 + 12.13 2.28 = 0.74 sm 5 O22 == 2227, 1.43 + 0.50 m 6* O39) = LAI 2.05 + 0.60 sm 7 74.03 + 8.03 2.10 = 0.32 sm 8 66.16 = 6.46 1.42 + 0.24 m 9 61.76 + 6.00 1.30 + 0.27 m 10 SI2D = O43 1.20 = 0.18 m 11 SIGS) 22 5.04! 1.90 + 0.15 sm 12 48.73 + 3.71 1.50 = 0.27 m 13 39)./D 22 Da) 1.40 + 0.12 m grandisonae and Diaz & Nufiez (1988) re- ported Alsodes verrucosus (adult and tad- poles). The other locality where Alsodes kaweshkari was collected is Seno Huemu- les. This area has the same ecological char- acteristics as Puerto Edén. The male para- type, collected at a border of a cold stream, also has the nuptial asperities on fingers and chest. Chromosomes.—Examination of 10 metaphase plates from the holotype re- vealed a diploid number of 2N = 26. All chromosomes are bi-armed and the funda- mental number (NF) is 52. Pairs 5, 8-10, 12, 13 are metacentric, pairs 1, 2, 4, 6, 7, 11 are submetacentric, and pair 3 is subtelo- centric. Pairs 1 and 4 have secondary con- strictions on the smaller arm, and pair 6 ex- hibits secondary constrictions on the longer arm. The karyotype of Alsodes kaweshkari is shown in Fig 5A. A summary of the rel- ative length, arm ratio, and type of chro- mosomes is presented in Table 2. The C-banded karyotype, based on five plates (Fig 5B) shows constitutive hetero- chromatin in the pericentromeric region of all chromosomes. Thin heterochromatic 527 bands can be discerned at some telomeres, especially in those of pairs 1, 2, 6 and 10. A thin insterticial band was observed in the long arm of pair 5. Pair 4 presents a poly- morphic situation: one chromosome shows a remarkable band of pericentromeric po- sition, but in the other the band is located in the centromeric region. This situation was observed in all the examined plates. Hemoglobin.—Figure 6A shows the chromatographic profiles of the hemoly- sates of Alsodes kaweshkari and A. monti- cola obtained from the fractions collected in the Sephadex G-50 minicolumn. The fractions with the higher absorbance values at 410 nm (fraction 9 in Alsodes kaweskari and fraction 8 in A. monticola) were elec- trophoretically analyzed under the experi- mental conditions used in this study. Figure 6B indicates the electrophoretical patterns obtained for these species. Both taxa share a similar two-banded pattern with an anodal band (Hb,) of 14.2 KDa an other catodal one (Hb,) of 16.8 KDa. Etymology.—The new taxon is named for the Kaweshkar indians, a brave people, hunters of sea-lions, who still live in Puerto Edén, the type locality of the new species. Comparisons When the adult snout—vent length of AI- sodes kaweshkari (males, 59.9—62.2 mm.) is compared with that of the other Alsodes species, it is observed that this frog is as large as A. nodosus (58.5 mm), A. barrioi (59.9 mm) and A. tumultuosus (61.5 mm). Other members of the genus are small [A. vittatus (37.4 mm), A. gargola (40.8 mm), and A. verrucosus (42.4 mm)] and medium- sized [A. monticola (49.1 mm), A. montan- us (50.2 mm), A. vanzolinii (51.1 mm), and A. pehuenche (52.3 mm)]. Among the large-sized frogs, the external morphology of the foot provides useful characteristics to identify each taxon. One example of this is A. nodosus, a species characterized by the absence of lateral fringes (present on the toes of A. barrioi, A. kaweshkari, and A. 528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A B 0.6 KDa 1 Js 3 0.5 A. kaweshkar! —-29.0— uy E 9414 monticola = , O < GS a 20.0— = OFZ 0.1 ~—_ ** Hb2 14.2— Hb; O O 4 8 12 16 20 Fractions Fig. 6. Hemoglobins of Alsodes kaweshkari and A. monticola. (A) Chromatographic elution profiles of A. monticola and A. kaweshkari in Sephadex G-50 minicolumn (B) Electrophoretical patterns of Alsodes kaweshkari and A. monticola. (1) Standard of molecular weight, (2) A. monticola, (3) Alsodes kaweshkari. tumultuosus). In the case of this latter spe- cies, its toes are long and thin, whilst in Alsodes kaweshkari and A. tumultuosus they are thick and strongly fringed. These frogs also differ in characteristics of the outer metatarsal tubercle (large in Alsodes kaweshkari and small in A. tumultuosus). On the other hand, both species differ in webbing; reduced in Alsodes kaweshkari and moderately developed in A. tumultuo- sus (Fig. 7A, B). Formas & Vera (1983) studied the kary- ological relationships among the member of the genus Alsodes. They recognized three groups within the genus: the barrioi group (2N = 34) (A. barrioi), the monticola group (2N = 26) (A. gargola, A. monticola, A. tumultuosus, A. vanzolinii, and A. verruco- sus), and the nodosus group (2N = 22) (A. nodosus). The presence of 26 chromosomes in Alsodes kaweshkari allows its inclusion in the monticola group. Muir (1981) studied the electrophoretical patterns of the hemoglobin molecule of 14 taxa of the frogs of the genus Xenopus. The species and subspecies were characterized and grouped according the number of bands obtained in the electrophoretical profiles. Xenopus borealis, X. fraseri, X. muelleri, X. ruwenzoriensis, X. tropicalis, X. vestitus and X. witteii show an electrophoretical pat- tern of two bands (Hb,, Hb,). Alsodes kaw- eshkari and A. monticola show a similar two-banded electrophoretical pattern like that found in some Xenopus species. Though our biochemical study was unable to differenciate Alsodes kaweshkari and A. monticola, this is the first attempt to char- VOLUME 111, NUMBER 3 529 A bas S = = 5mm Fig. 7. Schematic plantar views of the left foot of Alsodes kaweshkari (A) and A. tumultuosus (B). acterize the molecule of hemoglobin of de Chile (DBCG), Chile; Instituto de Biolo- frogs of the genus Alsodes. gia Animal, Universidad Nacional de Cuyo (IBA), Argentina; Instituto de Zoologia, Specimens Examined Universidad Austral de Chile (IZUA), Chile; Abbreviations.—Carmen Ubeda (personal Museo Nacional de Historia Natural collection) (CU), Argentina; Departamento (MNHN), Chile; Museo de Zoologia, Univ- de Biologia Celular y Genética, Universidad ersidad de Concepci6dn (MZUC), Chile. 530 Alsodes barrioi: YZUA 1629-1630; Cordil- lera Pelada, Provincia de Valdivia, 1020 m, Chile. Alsodes gargola: CU 6; Macizo Loncoluan, Provincia de Neuquén, 1900 m, Argen- tina. Alsodes montanus: IZUA 824; Estero Cob- arrubias, Provincia de Santiago, 2400 m, Chile. Alsodes monticola: TZUA 1550,1749; Cor- dillera Pelada, Provincia de Valdivia, 1020 m, Chile. Alsodes nodosus: YZUA 756,700; Aguas Claras, Provincia de Petorca, 150 m, Chile. Alsodes pehuenche: IBA 1643; Valle del Pehuenche, Provincia de Mendoza, 2500 m, Argentina. Alsodes tumultuosus: DBCG 161-162; La Parva, Provincia de Santiago, 2600 m, Chile. Alsodes vanzolinii: MZUC 12063-12070; Ramadillas, Provincia de Arauco, 100 m, Chile. Alsodes verrucosus: MNHN 1506; Puerto Edén, Provincia de Ultima Esperanza, 10 m, Chile. Alsodes vittatus: MZUC (untagged); Cor- dillera de Pemehue, Provincia de Malle- co) 1152) Chile: Acknowledgments The authors would like to give special thanks to Pablo Corti who collected the paratype in Seno Huemules. Alberto Velo- so, Patricia Iturra, José Navarro (Universi- dad de Chile), Enrique Pereira (Universidad Nacional de Cuyo, Argentina) and Carmen Ubeda (Universidad del Comahue, Argen- tina) kindly provided specimens for com- parisons. Marcos Navarro expertly prepared the drawings. This work was supported by Direccion de Investigacion Universidad Austral de Chile, Proyecto S 95-25. Literature Cited Bogart, J. P. 1970. Systematic problems in the am- phibian family Leptodactylidae (Anura) as in- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dicated by karyological analysis —Cytogenetics 9:369—-383. Cei, J. M. 1962. Batracios de Chile. Ediciones de la Universidad de Chile, Santiago, Chile, 128 pp. . 1980. Amphibians of Argentina.—Monitore Zoologico Italiano (N.S.) Monografia 2:1—609. Diaz, N., & H. Nunez. 1988. Nuevo hallazgo de Al- sodes verrucosus (Philippi 1902) en Chile y descripciédn de su larva.—Boletin Museo Na- cional Historia Natural, Chile 41:87—94. di Castri, EF 1968. Esquisse écologique du Chili. Biol- ogie de l’Amerique australe. Cl. Debouteville and E. Rappaport, eds.—Editions du Centre Na- tional de la Recherche Scientifique, Paris 1V:7— 2. Duellman, W. E. 1970. The hylid frogs of the Middle America.——Museum of Natural History, Uni- versity of Kansas Monographs 1:1—753. Formas, J. R. 1989. Sinonimia e identidad de la rana austral Chilena Eupsophus vittatus (Philippi 1902) (Anura: Leptodactylidae).—Boletin So- ciedad de Biologia, Concepci6n 60:123—127. . 1991. The karyotypes of the Chilean frogs Eupsophus emiliopugini and E. vertebralis (Amphibia: Anura: Leptodactylidae).—Pro- ceedings of the Biological Society of Washing- ton 104:7-11. , & M. I. Vera. 1983. Karyological relation- ships among frogs of the genus Alsodes, with description of the karyotypes of A. vanzolinii and A. verrucosus.—Copeia 1983:1104—1107. Frost, D. R. 1985. (ed). Amphibian species of the world. A taxonomic and geographic reference. Allen Press and the Association of Systematics Collection, Lawrence, Kansas, 732 pp. Laemmli, U. K. 1970. Cleavage of structural proteins during the assambly of the head of bacterio- phage T,.—Nature 227:680—685. Levan A., Fredga A., & A. Sandberg. 1964. Nomen- clature for centromeric positions on chromo- somes.—Hereditas 52:201—220. Lynch, J. D. 1975. A new Chilean frog of the extra- andean assamblage of Telmatobius (Amphibia: Leptodactylidae).—Bulletin of Southern Cali- fornia Academy of Sciences 74:160—161. Muir, A. V. 1981. Comparison of hemoglobins from the genus Xenopus (Amphibia: Salientia).— Journal of Experimental Zoology 218:327-—333. Philippi, R. A. 1902. Suplemento de los Batraquios Chilenos descritos en la Historia Fisica y Poli- tica de Chile de don Claudio Gay. Santiago, 61 Pp. Sumner, A. T. 1972. A simple technique for demon- strating centromeric heterochromatin.—Experi- mental Cell Research 75:304—306. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):531—534. 1998. Helicoprion nevadensis (Wheeler, 1939) from the Pennsylvanian—Permian Antler Peak Limestone, Lander County, Nevada (Pisces: Selachii: Helicoprionidae) Rex Alan Hanger and Ellen E. Strong (RAH) Department of Geology, George Washington University, Washington, D.C. 20052, U.S.A.; (EES) Department of Biological Sciences, George Washington University, Washington, D.C. 20052, U.S.A. Abstract.—A new specimen of Helicoprion nevadensis (Wheeler 1939) is described from the Antler Peak Limestone of Lander County, Nevada. The occurrence is significant as the only other individual of the species known has no geographic or stratigraphic information. The new specimen is dated as Wolf- campian (Early Permian) using associated fusulinids. Presence of H. nevadensis in Nevada and similar species in California confirms open-marine connections between the Permian shelf of North America and coeval island arcs to the west. A new specimen of the spiral tooth-whorl shark fossil, Helicoprion nevadensis (Whee- ler 1939) is described from the Pennsylva- nian-Permian Antler Peak Limestone of Lander County, Nevada. This is only the second individual of the species discovered. The original geographic and stratigraphic lo- cation of the holotype are unknown. In con- trast, this specimen is precisely located and accurately dated for the first time. It is Wolf- campian (Lower Permian) based upon co- occurrence of fusulinids. Although this oc- currence does not resolve the controversy surrounding longitudinal separation between the North American continental margin and coeval island arcs to the west (Harwood and Miller 1990), this accurate locality and bio- Stratigraphic information does allows us to conclude that open marine connections ex- isted between North American (autochtho- nous) shelf seas and island arc (allochtho- nous) deposits during the Early Permian. Systematic Paleontology Family Helicoprionidae Bendix-Almgreen, 1966 Genus Helicoprion Karpinsky, 1899 Helicoprion nevadensis Wheeler, 1939 Fig.) 1 Helicoprion nevadensis.—Wheeler, 1939: 109-112, fig. 3. Description.—The symphysial whorl of the specimen consists of 1.5 volutions and is broken off at both juvenile and adult ends (Fig. 1). The maximum preserved diameter (tooth crown tip to tooth crown tip) is 100.35 mm. Description is compromised by the fact that the enclosing rock is broken so that most of the specimen is split laterally, and several tooth crowns are broken off at the shaft and missing. 47 tooth crowns are preserved in one volution. For a well-pre- served tooth at approximately 1 volution, the volution height (V) is 20.2 mm, the shaft height (S) is 2.3 mm, with an S/V ra- tio of 0.11. No serration denticles are pre- served. Depository.—The specimen is deposited in the University of California Museum of Paleontology (UCMP) as specimen 140632. Location.—From UCMP Locality V94012, in western Lander County, Neva- da, with a Township/Range coordinate of T32N, R44E, in NE % of SE % of Section 21 on the Snow Gulch, Nevada, 7.5-minute U.S. Geological Survey quadrangle, 1991 edition. The specimen occurs in pale brown PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. X 1.1. siltite of the siliclastite-dominated facies of the Antler Peak Limestone (Theodore 1994). Comparison.—Several workers (Bendix- Almgreen 1966, Siedlecki 1970) have noted the very strong similarities of all described Helicoprion species. Unfortunately, Heli- coprion fossils are always found as isolated specimens, not populations. The possibility exists that much of the published species diversity of the genus actually reflects the range of morphologic variation of very few, or even a single species. Because of this taxonomic imprecision and the incomplete- ness of UCMP 140632, specific taxonomic placement is difficult. For S/V ratio and number of tooth crowns/volution, UCMP 140632 is most like Wheeler’s (1939) ho- lotype H. nevadensis, UCMP 1001. Helicoprion nevadensis. (UCMP 140632) from the Antler Peak Limestone, Lander County, Nevada, Biostratigraphy.—UCMP Locality V94012 occurs between strata containing fusulinid foraminifera (Verville et al. 1986). Several species of Schwagerina confirm a Wolfcampian age for UCMP 140632, con- sistent with the established geologic range of Helicoprion (Wolfcampian—Guadalupi- an). Biogeography.—The genus Helicoprion has a worldwide distribution, with species described from Russia (Karpinsky 1899, Obruchev 1953, for summary), Japan (Yabe 1903, Araki 1980), Laos (Hoffet 1933), Australia (Teichert 1940), Spitsbergen (Siedlecki 1970), Arctic Canada (Nassichuk & Spinosa 1970, Nassichuk 1971), British Columbia, Alberta (Logan & McGugan 1968), Idaho, Wyoming (Hay 1907, 1909; Williams & Dunkle 1948; Bendix-Alm- VOLUME 111, NUMBER 3 green 1966), Nevada, California (Wheeler 1939, Larson & Scott 1955, this report) West Texas (Kelly & Zangerl 1976, Chorn 1978), and Mexico (Mulleried 1940), Thus, Helicoprion is found within rocks of many Permian benthic faunal provinces (see for example, Shi et al. 1995, Yancey 1975, Bambach 1990). Collectively, these prov- inces span polar to equatorial paleolatitudes in both hemispheres, a remarkable distri- bution, even for a nektic organism, sug- gesting that temperature was not a limiting factor for Helicoprion. Paleogeography.—Documenting the oc- currence of Helicoprion is problematic in the paleogeographically “‘suspect’’ accreted terrane region of western North America. Besides UCMP 140632, only three individ- ual specimens are known. The holotype of H. nevadensis Wheeler, 1939, described originally as from the Koipato Group of the Humboldt Range, Nevada, is considered by Silberling (1973) not to have been obtained from the Koipato Group because of dis- crepancies between the lithology of the rock fragments attached to the fossil and the diagnostic lithologies of the Koipato Group. Helicoprion sierraensis Wheeler, 1939 is said to be found by an amateur “‘in a gla- cially transported boulder in the valley of Frazier Creek, California.’’ Convoluted in- terpretations have to be proposed to attempt to determine provenance for the fossil. A third reported specimen from Nevada, H. sp., is reasonably located, but comes from autochthonous rocks near Elko, Nevada (Larson & Scott 1955). This specimen is not figured in any publication, nor is its mu- seum deposition mentioned. The current lo- cation of the specimen is unknown. Heretofore, lack of adequate documen- tation for the genus has prevented reliable paleogeographic inference. The joint occur- rence of Helicoprion in autochthonous rocks of the Early Permian continental mar- gin (Wheeler 1939, Larson & Scott 1955, Roberts et al. 1958, this report) and in the allochthonous Northern Sierra terrane of McCloud Belt faunal affinity (Harwood 533 1992, Miller 1987) should not be taken as evidence of original paleogeographic prox- imity of the McCloud Belt to North Amer- ica during the Permian, nor can it resolve the problem of longitudinal separation. However, presence of the nektic genus in both areas does suggest presence of open marine conditions between these tectonic elements. Acknowledgments R. T. Ashinhurst of the Battle Mountain Gold Company, Battle Mountain, Nevada kindly donated the specimen to UCMP. Literature Cited Araki, H. 1980. The discovery of the fossil genus Hel- icoprion, a chondrichthian, from Kesennuma City, Miyagi Prefecture, Japan.—Geological Society of Japan, Journal 86:135—-137. Bambach, R. K. 1990. Late Paleozoic provinciality in the marine realm. Pp. 307-323 in W. S. Mc- Kerrow and C. R. Scotese, eds., Palaeozoic Pa- laeogeography and Biogeography. Geological Society of London Memoir Number 12, Lon- don, 431 pp. Chorn, J. 1978. Helicoprion (Elasmobranchii, Edesti- dae) from the Bone Spring Formation (Lower Permian) of West Texas.—University of Kansas Paleontological Contributions, Paper 89:2—4. Bendix-Almgreen, S. E. 1966. New investigations on Helicoprion from the Phosphoria Formation of South-East Idaho, U.S.A.—Biologiske Skrifter Danske Videnskabermes Selskab 14(5):1—54. Harwood, D. S. 1992. Stratigraphy of Paleozoic and Lower Mesozoic rocks in the Northern Sierra terrane, California.—United States Geological Survey Bulletin 1957:1—78. , & M. M. Miller (eds.). 1990. Paleozoic and Early Mesozoic Paleogeographic Relations; Si- erra Nevada, Klamath Mountains, and Related Terranes. Geological Society of America, Spe- cial Paper 255, Boulder, Colorado, 422 pp. Hay, O. P. 1907. A new genus and species of fossil shark related to Edestus Leidy.—Science, New Series 26(653):22—24. . 1909. On the nature of Edestus and related genera, with descriptions of one new genus and three new species.—Proceedings of the U.S. National Museum 37:43—61. Hoffet, J. H. 1933. Etudes géologiques sur le contre de Il’Indochine entre Tourane et le Mékong.— Bulletin, Service Géologique de 1’Indochine 20(2):1-154. 534 Karpinsky, A. 1899. Uber die Reste von Edestiden und die neue Gattung Helicoprion.—Verhandlung Russisch-Keiserling Mineralogische Gesells- chaft zu St. Petersburg Series 2, 36(II]):361—376. Kelly, M. A., & R. Zangerl. 1976. Helicoprion (Edes- tidae) in the Permian of West Texas.—Journal of Paleontology 50:992—994. Larson, E. R., & J. B. Scott. 1955. Helicoprion from Elko County, Nevada—Journal of Paleontolo- gy 29:918-919. Logan, A., & A. McGugan. 1968. Biostratigraphy and faunas of the Permian Ishbel Group, Canadian Rocky Mountains.—Journal of Paleontology 42:1123-1139. Miller, M. M. 1987. Dispersed remenants of a north- east Pacific fringing arc: Upper Paleozoic island arc terranes of McCloud Belt faunal affinity, western United States.—Tectonics 6:807—830. Mullereid, EF K. G. 1945. El edestido, Helicoprion, en contrado por primera vez en Mexico, en el es- tado de Coahuila.—Ciencia Revista Hispano- America de Ciencias Puras y Aplicadas 6:208— 211. Nassichuk, W. W. 1971. Helicoprion and Physonemus, Permian vertebrates from the Assistance For- mation, Canadian Arctic Archipelago—Geo- logical Survey of Canada, Bulletin 192:83—93. , & C. Spinosa. 1970. Helicoprion sp., A Perm- ian elasmobranch from Ellesmere Island, Ca- nadian Arctic.—Journal of Paleontology 44: 1130-1132. Obruchev, D. V. 1953. A study of the edestids and the work of A. P. Karpinsky.—Akademia Nauk SSSR, Trudy Paleontologischkogo Instituto 45: 1-88. Roberts, R. J., PR E. Hotz, J. Gilluly, & H. G. Ferguson. 1958. Paleozoic rocks of North-Central Neva- da.—American Association of Petroleum Ge- ologists Bulletin, 42:2813—2857. Shi, G. R., N. W. Archbold, & L.-P. Zhan. 1995. Dis- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tribution and characteristics of mixed (transi- tional) mid-Permian (Late Artinskian—Ufimian) marine faunas in Asia and their palaeogeo- graphical implications.—Palaeogeography, Pa- laeoclimatology, Palaeoecology 114:241—271. Siedlecki, S. 1970. A Helicoprion from the Permian of Spitsbergen.—Norsk Polarinstitutt Arbok 1968:36—54. Silberling, N. J. 1973. Geologic events during Perm- ian-Triassic time along the Pacific margin of the United States, Pp. 345-362 in A. Logan and L. V. Hillis, eds., The Permian and Triassic sys- tems and their mutual boundary. Canadian So- ciety of Petroleum Geologists, Memoir 2. Cal- gary, Alberta., 766 pp. Teichert, C. 1940. Helicoprion in the Permian of West- ern Australia—Journal of Paleontology 14: 140-149. Theodore, T. G. 1994. Preliminary geologic map of the Snow Gulch quadrangle, Humboldt and Lander Counties, Nevada. United States Geological Survey, Open-file Report 94-436. Verville, G. J.. G. A. Sanderson, & D. D. Drowley. 1986. Wolfcampian fusulinids from the Antler Peak Limestone, Battle Mountain, Lander County, Nevada.—Journal of Foraminiferal Re- search 16:353-—362. Wheeler, H. E. 1939. Helicoprion in the Anthracolithic (Late Paleozoic) of Nevada and California, and its stratigraphic significance.—Journal of Pale- ontology 13:103—114. Williams, J. S., & D. H. Dunkle. 1948. Helicoprion- like fossils in the Phosphoria Formation.—Geo- logical Society of America, Bulletin 59:1362. Yabe, H. 1903. On a Fusulina-Limestone with Heli- coprion in Japan.—Geological Society of To- kyo, Journal 10:1—13. Yancey, T. E. 1975. Permian marine biotic provinces in North America.—Journal of Paleontology 49: 758-766. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):535-—550. 1998. Pentamera rigida and P. pediparva, two new species of sea cucumber from the west coast of North America (Echinodermata: Holothuroidea) Philip Lambert Natural History Section, Royal British Columbia Museum, P.O. Box 9815 Stn Prov Govt, Victoria, B.C., Canada V8W 9W2 Abstract.—Two new species of the genus Pentamera from the west coast of North America are described, and all known species from the west coast of North and South America are reviewed. Pentamera rigida is a slender, stiff, U-shaped species with five rows of podia. It occurs in sand-gravel substrata from 18 to 421 m between British Columbia and California. The table ossicles of the skin have an angular disc with a broad, low, convoluted spire. Pentamera pediparva has a soft, curved body with five rows of fine podia. It is known from central British Columbia to northern California from 8 to 120 m in mud- gravel or sand-gravel. The table ossicles are small and oval with small, two pillared spires. The taxonomic characters of fourteen species of Pentamera are summarized. Twelve species of Pentamera have been described from the west coast of North and South America (Stimpson 1851, 1864; Lud- wig 1886a, 1886b; Clark 1924; Deichmann 1938a, 1938b). Panning (1949) placed the genus Pentamera in Thyoninae, a new sub- family of Cucumariidae. Based on the pres- ence of posterior processes composed of a mosaic of small pieces, Pawson & Fell (1965) transferred Thyoninae Panning to the family Phyllophoridae. No new species of Pentamera have been added to the west coast fauna since Deichmann (1938a, 1938b), with the exception of those trans- ferred to the genus by Panning (1949). The purpose of this paper is to describe two new species of Pentamera from the Pacific coast of North America and present a summary table of morphological characters for spe- cies from the west coast of North and South America. Materials and Methods Ossicle slides were prepared by the method described in Lambert (1985). At least 30 ossicles per specimen were mea- sured on transects across each slide. Length and width of disc, and width of central spire were recorded for tables. Length, height of curve, and height of spire were recorded for supporting tables (Fig. 7). Only ossicles that were totally within the field of vision, lying flat, and not broken, were measured. Via a drawing tube attached to a Wild M20 microscope, the image of each ossicle was visualised on a digitising tablet (Summas- ketch II, Summagraphics Corporation). Measurements were made with the cursor and automatically saved to a file using SigmaScan software (Jandel Scientific). That file was then transferred to a statistical package (Systat, Systat, Inc.) for analysis. External measurements of the whole spec- imen were made with an ocular micrometer, calipers, or in the case of strongly curved specimens, a string was used to trace the distance from the base of the tentacles to the anus, around the outer side of the curve, usually the ventral side. Institutions mentioned in this paper are abbreviated as follows: Allan Hancock 536 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. length 7 cm, RBCM 984-199-3. Foundation (AHF) now housed at LACM; California Academy of Sciences, San Fran- cisco (CASIZ); Fisheries Research Board of Canada (FRB) now known as Department of Fisheries and Oceans, (DFO); Canadian Museum of Nature, Ottawa (CMNI or NMC); Natural History Museum of Los Angeles County (LACM); Royal British Columbia Museum, Victoria (RBCM); Na- tional Museum of Natural History (USNM), Washington, D.C. Results Order Dendrochirotida Grube Family Phyllophoridae Oestergren Diagnosis.—Dendrochirotid holothurians without a test. Tentacles 10—25 (usually 10 or 20), well branched. Pedicels either re- stricted to the ambulacra or scattered, but fewer and usually papilliform dorsally. Cal- careous ring complex, often tubular, with long or short posterior processes, both ring and processes always composed of a mo- saic of small pieces. Ossicles usually plates or buttons, rods, tables or derivatives of ta- bles (after Thandar 1990). Remarks.—Thandar modified the diag- nosis of Pawson (1982) by adding that the podia may be in rows or scattered and that Preserved specimen of Pentamera rigida dredged from type locality, 115 m, off Nootka Sound, B.C., the ossicles may be tables or derivatives of tables in the form of plates, rods or buttons. Subfamily Thyoninae Panning, 1949 Diagnosis.—Small to medium-sized phyllophorids, rarely more than 100 mm long. Body soft. Pedicels numerous, usually scattered all round, but often most crowded ventrally. Tentacles 10, ventral two always reduced. Calcareous ring tubular with long paired posterior processes on radial plates. Ossicles of body wall usually in the form of tables or plates (buttons), or absent (after Thandar 1990). Genus Pentamera Ayres, 1852 Diagnosis.—Small to medium sized forms. Ten tentacles with two smaller ven- tral tentacles. Nonretractile podia in five se- ries, not scattered. Radials of the calcareous ring with long forked tails. In the skin two pillared tables or derivatives of these with the spire reduced or absent. Podia with large endplates and curved supporting ta- bles with spires varying from low to high. Tentacles usually with rods, plates or both. (After Deichmann 1941) Type species: Pentamera pulcherrima Ayres VOLUME 111, NUMBER 3 33) Fig. 2. introvert. (C) Tentacle ossicles. (D) Supporting tables and end plates of podia. (E) Calcareous ring and retractor muscles; scale bar 5 mm. All from holotype (RBCM 997-123-1) except for the podial end plates (RBCM 983- 1658-30) and the calcareous ring of paratype (RBCM 984-199-3). Top scale bar 100 ym applies to all ossicles. Pentamera rigida, new species Figs. 1-3 Pentamera lissoplaca.—Bergen 1996 (par- tim):239, fig. 9.23B. Specimens de- scribed as having “‘large tables’’ may be P. rigida. Pentamera sp. A Lambert, 1997:101, figs. 52-53, photo 24. Diagnosis.—Body U-shaped; skin usu- Ossicles and calcareous ring of Pentamera rigida. (A) Tables of the dorsal skin. (B) Plates from the ally stiff and white. Podia in five series, longer and more abundant in midventral re- gion than distally. Ten small dendritic ten- tacles including smaller ventral pair; often speckled with brown. Calcareous ring long and tubular with forked tails on the radials; made up of a mosaic of smaller pieces; ratio of radial to interradial length approximately 4:1. Skin ossicles circular, triangular or star- shaped tables (100-300 sm) with a broad 538 Fig. 3. Collection sites for Pentamera rigida. convoluted spire covering about one-half width of disk; small, curved supporting ta- bles (100 pm) with low spire. Introvert with oval knobbed perforated plates, some with remnant of a spire. Tentacle ossicles in two forms, finer curved oval plates and large ro- bust rods with a few holes. Material examined.—Eighty-one speci- mens from 15 localities in British Colum- bia, one locality in Oregon, four in Califor- nia, and one from an unknown locality on the west coast of U.S.A. Number of speci- mens in parentheses after the catalogue number; depth in metres (m). Holotype.—RBCM 997-123-1 (1), col- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lected by E Bernard and T. C. Lambert, FRB station 63-91, 8 Sep 1964, length 6.8 cm (measured along the outside of the curved body), female. Type locality.—British Columbia, Van- couver Island, Nootka Sound, 49°32'N, 127°03'W, 115 m, sandy mud. Paratypes.—RBCM 984-248-1 (1), col- lected by D. B. Quayle, FRB Haul No. 1, 7 Jul 1962, British Columbia, Kitkatla Inlet, Porcher Island, 53°51'N, 130°W, 37 m, sand and gravel, specimen 10.8 cm long. RBCM 984-200-1 (7), FRB station 63-204, 10 Sep 1964, British Columbia, Vancouver Island, Quatsino Sound, 50°19.6’N, 128°06.5'W, 119 m, sand, specimen lengths from 2.8 to 5.1 cm. RBCM 984-199-3 (12), collected by E Bernard and T. C. Lambert, FRB sta- tion 63-91, 8 Sep 1964, British Columbia, Vancouver Island, Nootka Sound, 49°32’N, 127°03'W, 115 m, sandy mud, from same lot as holotype, specimen lengths from 3 to 8 cm (mean 5.5). USNM E16915 (32), col- lected by A. Carey, Oregon State Univer- sity, Cruise C 700810, BMT 239, 17 Aug 1970, Oregon, off Hecata Head Light, 44°08.8'N, 124°24'W, 100 m, specimen lengths from 1.5 to 5.7 cm. CASIZ 50227 (1), Sta. 20, 28 Sep 1949, California, off Point Reyes, south of Cordell Bank, 37°55.75'N; 123°19:25'W,. PLO= ise specimen 6 cm long. AHF 440.61 (1), Ve- lero 6131-59, California, Ventura Co., Ox- nard, Port Hueneme Lighthouse, 34°8.1'N, 119°21.5'W, 165 m. Other collections.—British Columbia: RBCM 989-564-4 (1), collected by D. B. Quayle, FRB station 70-24, Jul 1970, Bramham Island near Pine Island, 51°05’N, 27°39'W, 82 m. RBCM 984-213-1 (1), col- lected by D. B. Quayle, FRB station 71-14, Apr 1971, Gordon Channel, near Nigei Is- land, 50°53. 1'N,: 127°36.3' W942 See RBCM 990-939-11 (2), FRB station 67-66, Aug 1967, Vancouver Island, Cape Scott, 50°39.8'N, 128°47.2'’W, 200 m, gravel. RBCM 984-249-1 (1), collected by J. Fleu- ry, FRB Drag #7 + #8, 27 Jun 1962, Baker Pass, near Cortes Island, 50°04.0’N, VOLUME 111, NUMBER 3 539 Fig. 4. Holotype of Pentamera pediparva collected by scuba from 7.5 m in Quatsino Sound, B.C., length 8 cm, RBCM 997-124-1. 124°59.0’W, 117 m, gravel. RBCM 980- 344-1 (1), collected by P. Lambert, station L80-61, 6 Jul 1980, Vancouver Island, Kyu- quot Channel, Sandy Bay, Rugged Point, 49°58.2’N, 127°14.6'W, 9 m, clean sand and algae. RBCM 55-56 (1), collected by D. El- lis, 16 Aug 1965, Strait of Georgia, 49°54.05'N, 125°04’W, 178 m, sand. RBCM 983-1658-30 (3), collected by Gor- don Green, 23 Nov 1983, Vancouver Island, off Kyuquot Sound, 49°45.7'N, 127°30.1’W, 150 m, sand. RBCM 974-570- 6 (3), FRB station 2231-33, 17 Jul 1934, Vancouver Island, Estevan Point, 49°22.3'’N, 126°55'W, 137 m, sand. RBCM 986-93-27 (3), collected by P. Lambert, on Endeavour, Station L86-9, 19 Mar 1986, Vancouver Island, Estevan Point, 49°11'N, 126°45.2'W, 120 m, sand. RBCM 983- 1397-1 (1), collected by M. Byrne, 11 Feb 1983, east of Galiano Island, 49°01.48'N, 123°29.45'W, 50—230 m. RBCM 991-10-1 (2), collected by S. Carson, 22 Feb 1990, Barkley Sound, Trevor Channel, 48°52'N, 125°08'W, 54 m, mud. RBCM 988-758-10 (1), collected by P. Lambert, G. Green, D. Bright, Station L88-27, 22 Jun 1988, Sat- ellite Channel, Boatswain Bank, 48°42.2'N, 123°32.1'W, 18 m. California: CASIZ uncatalogued (1), 9 Aug 1940, 2.5 mi. off Mad River, Eureka, 37 m. USNM E2371 (4), Albatross St. 2902, California, Channel Islands, Santa Cruz Channel, Santa Rosa Island, 34°6.0'N, 120°2.0'W, 97 m. USNM 30563 (1), Alba- tross St. unknown, West Coast United States, depth unknown. Description.—Total length 1.5-10.5 cm (measured along the outer curve of the body); mean length (cm + 1 SD) 4.6 + 1.6 cm (n = 72). Holotype 6.8 cm long. Body typically bent in a tight U-shape with long slender anterior and posterior ends (Fig. 1). Skin stiff with ossicles. Podia slender and probably non-retractile because of their heavy complement of ossicles. Podia form five bands, each consisting of two rows, crowed together in the middle of the body on ventral side but sparse distally and on dorsal side. Specimens typically white in life and in alcohol. Ten tentacles arranged in five pairs with a smaller ventral pair. Madreporite in dorsal mesentery about two-thirds of distance from anterior end of calcareous ring. Semi-circular madreporite with a long narrow stone canal, connects to the ring canal near posterior tip of calcar- eous ring. One polian vesicle usually on ventral side of ring. Two respiratory trees emerge from cloa- ca on left and right sides; each one splits into a dorsal and ventral branch; dorsal Fig. 5. the introvert. (C) End plate of podia (D) Tentacle ossicles. (E) Supporting tables of podia (F) Calcareous ring; scale bar, 5 mm. All from holotype (RBCM 997-124-1). Top scale bar (100 tm) applies to all ossicles. branch runs full length of body, ventral branch only to mid body. Respiratory trees usually brown, with a main trunk and sim- ple, short side branches. Cloaca occupies most of narrow ‘“‘tail.”’ Gonad has two tufts of unbranched tu- bules, one tuft on each side of dorsal mes- entery where it joins the dorsal body wall. Thin retractor muscles attach to body wall at a point level with posterior end of calcareous ring. Five thin longitudinal mus- cles. Radial plates of calcareous ring with long posterior prolongations; interradials with no tails, shaped like an elongate tri- angle (Fig. 2E). Transverse thickenings in- dicate joints between mosaic of pieces that make up the tails. Ratio of interradial to ra- dial length about 1:4. Tables of mid skin circular to roughly tri- See ° PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ossicles and calcareous ring of Pentamera pediparva. (A) Tables of the dorsal skin. (B) Plates from angular or star-shaped, with a mean diam- eter (um + 1 SD) of 194 + 45 (n = 141) with a complex central spire (width 102 + 32, n = 140) that covers about one-half the surface area of table (Fig. 2A). Supporting tables of podia small (length 110 + 15,n = 90) and tightly curved with a low spire (height 16 + 5, n = 90) with three or four teeth. Ratio of length (L) to total height (TH) of supporting tables 1.9 + 0.3, n = 90 (see Fig. 7 for explanation of dimen- sions). Podial end plates small (diameter 115-135 pm, n = 5) (Fig. 2D). Ossicles in tail region not significantly different from mid skin. In juveniles (<2 cm) tables tend to be more star-shaped and spires not as ro- bust. Introvert has oval perforated plates with two larger central holes and knobs on the surface of plate (Fig. 2B). In tentacles, VOLUME 111, NUMBER 3 Fig. 6. Collection sites for Pentamera pediparva. ossicles range in size from large flat rods (ength 230 wm) with a few knobs and holes to smaller curved perforated plates with scalloped edges (Fig. 2C). Etymology.—The species name, rigida, is based on the Latin word rigidus, meaning stiff or inflexible and refers to the typically stiff ossicle-filled body of this species. Distribution and habitat.—Pentamera rigida is known from Porcher Island near Prince Rupert, British Columbia (53°51'N, 130°W) south to Santa Cruz Channel, Santa Rosa Island, Channel Islands, California (34°6.0'N, 120°2.0’W) (Fig. 3). It ranges in depth from 18 to 421 m on the continental 541 = IOS x shelf. The majority of collections are from less than 200 m. The one specimen from 421 m is from a deep trench between Van- couver Island and the mainland. So far none have been collected on the continental slope or deeper. Usually occurs in sand or gravel substrata. Pentamera pediparva, new species Figs. 4-6 Pentamera sp. B Lambert, 1997:104, figs. 54, 55, photo 25. Diagnosis.—Body curved; up to 7 cm long; skin usually soft; colour white to 542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Pentamera of the west coast of North and South America (ossicle measurements in micrometers + 1 SD; L = length, H = height of spire, TH = total height, n = 15 unless otherwise indicated). Pentamera species Distribution Body shape Podia Colour P. beebei Deichmann 1938 Costa Rica Small form Typical White P. calcigera (Stimpson Arctic; eastern Russia; Curved with tapered Single pairs in White or yel- 1851) as Pentacta Alaska, Bering Sea, posterior; 50-60 5 rows; non- lowish P. charlottae Deichmann 1938 P. chierchia (Ludwig 1886) as Cucumaria P. chiloensis (Ludwig 1886) as Cucumaria P. lissoplaca (Clark 1924) as Cucumaria P. montereyensis Deich- mann 1938 Chukchi; 7—25 fm: 33-80 m Monterey Bay to southern California (Bergen); intertidal to 642 m Panamic region; Mex- ico and Costa Rica; shallow water Cape Hom to Lower California; 8-60 fm (Deichmann) Southeast Alaska south to Baja Cali- fornia; 9-82 m Pacific Grove, Long Beach & Santa Rosa I. California, intertidal & shallow on rocks mm Few cm long; body straight. Up to 6.5 cm, cylindrical (Bergen) Small 3—6 cm; soft- skinned Curved, about 5 cm; skin thin and wrin- kled Curved, tapering to posterior Moderate-size form, up to 5 cm; cylin- drical retractible 5 double rows; cylindrical, non-retractile Numerous cy- lindrical feet in 5 bands “‘Resembles ge- nus type’; conspicuous cylindrical podia in 5 well-defined bands 5 crowded dou- ble series; non-retractile stiff with os- sicles 5 bands, cylin- drical, non- retractile White or cof- fee (pre- served) Dark brown almost black. White Yellowish white in preservative White, tenta- cles and in- trovert with few pigment spots VOLUME 111, NUMBER 3 Table 1.—Extended. 543 Calcareous ring Mid-dorsal Ossicles Introvert Tentacles Not stated in origi- nal description Radial with long posterior pro- cesses; shorter inter-radial Radials with long posterior pro- cesses; interradi- al % length, no tails Long posterior pro- cesses Not stated in origi- nal description Ring rather high; interradials about 2 mm high and half as wide concave behind, radials with pos- terior processes. Interradial to ra- dial 1:3 ~ Minute tables (L = 37, n = 2) with four+ holes, 2-pillared spire with tuft of spines; curved support tables (L = 65 + 9, n = 6) with tall spire, ratio of L to TH 0.8 + 0.3; large endplate. Data from illustrations. Mostly oval plates (L = 162 + 35); fewer oval or squarish tables with central spire. Curved supporting ta- bles (L = 163 + 20) with moderate spire (H = 36 = 6), ratio of L to TH = 1.8 + 0.3 Well developed end plates with indented perimeter (136-177, n = 7). Small oval tables (L = 91 + 9) with 2-pillared spire. Curved supporting tables similar to tables but with a taller spire (H = 59 + 9), having 2 or 3 crossbeams and topped with a few teeth. Large end-plates (130-— 225, n = 7). Ossicles vary in number but not crowded; tables (L = 70, n = 3) with 4 central holes and dentate margin, low 2 pillared spire often lacking; few spectacle rods, trace of spire; podia with large end-plate. Tables with oval to squarish disk (L = 69 + 13, n = 6) with 4 or 8 holes; 2 pillared spire, and 1 or 2 crossbeams short teeth on top; po- dia with large end-plate & curved supporting tables (L = 90 + 105, n = 2) with low spire few flat teeth; ratio of L to TH = 1.5; data de- rived from illustrations of ossicles. Ossicles in two layers; scattered deli- cate tables (diameter 30—50) with fine spire; inner layer smooth, crowded, diamond-shaped plates (L = 120 + 15); curved supporting ta- bles (L = 103 + 10) with moderate spire (H = 36 + 6), ratio of L to TH = 1.9 + 0.3; podia end plates 90-110, n = 4. Small oval buttons (diameter 25—55 jum) with 2 central and up to 8 marginal knobs; supporting tables with marginal knobs, reduced spire; well developed end-plates (235— 275, n = 4), perforated rods. Unknown Oval perforated plates with meshwork at center (this study) Tables with oblong disk, dentate margin and low spire No specimen exam- ined; introvert un- known Rosettes Oval perforated plates (100-200) with 2 larger central and numerous smaller holes, many knobs on surface rated plates with 2 larger central holes and 6—20 other holes; 2 central knobs As for genus Ten; 8:2 small Ten (8:2); with rods, plates, and tables Ten (8:2); curved per- forated plates and rods Tentacles missing in type, Deichmann— typical for genus; with delicate rods Ten (8:2); ossicles of two classes: large (~300) robust perfo- rated rods and deli- cate (~100) oval perforated plates with 2 large central holes, wavy edge Round to oblong perfo- Ten (8:2) 544 Table 1.—Continued. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pentamera species Distribution Body shape Podia Colour P. populifera (Stimpson 1864) as Pentacta P. pseudocalcigera Deich- mann 1938 P. pseudopopulifera Deichmann 1938 P. trachyplaca (Clark 1924) as Cucumaria P. zacae Deichmann 1938 P. rigida, new species Kodiak I. to southern California perhaps Baja; to 60 fm Southeast Alaska to San Benitos Island, Baja California, 7— 250 fm Southern California; shallow water to 57 fm (Bergen) Queen Charlottes to Channel Is, Califor- nia, to 33 m Tangola Bay, Mexico; 23 fm Porcher I., B.C. south to Santa Rosa L., Channel Islands, California Thick fusiform up to 6 cm, skin thin and flexible; curved with tapering poste- rior or nipple-like Medium sized, U- shaped, tapering to blunt ends, rather stiff Medium size to 6.5 cm variable fusi- form curved and skin usually wrin- kled Small cylindrical, up to 3 cm; blunt sub- equal ends Small (few cm) with strongly curved body tapering to- ward ends U-shaped, stiff, elon- gate, fatter in mid- dle and tapering at both ends, posterior more so 5 bands small numerous po- dia full length of body In 5 raised bands, podia conical in shape, non- retractile 5 crowded bands of po- dia, up to 8 across; non- retractile; cylindrical 5 bands stout podia Numerous cy- lindrical, non- retractile po- dia in 5 bands, more on ventrum than at oral or anal ends 5 bands fine, stiff podia, usually more abundant midventrally Creamy white Podia usually lighter than the pinkish inter ambu- lacrum Straw-colored podia, beige in alcohol Yellowish white in al- cohol, tenta- cles darker Dirty white Whitish VOLUME 111, NUMBER 3 Table 1.—Continued. Extended. Calcareous ring Radials with long posterior pro- cesses about twice length of interradials Long posterior pro- cesses Radials with long posterior pro- cesses, radial about 3 times longer than in- terradial Posterior processes of medium length, interradi- als an elongate triangle No calcareous ring in type Typical Pentamera with very long posterior pro- cesses; approxi- mately 1:4 Mid-dorsal Ossicles Oval to star-shaped tables (L = 213 + 58) a few reduced in size, with scalloped edge; 25—40 holes; nar- row central spire (width at base 48 + 14); curved supporting tables (L = 119 + 14) with a tall spire (H = 41 + 7) topped with teeth, ratio of L to TH = 1.4 + 0.2; podia have smallish endplates with notched pe- rimeter (90-105, n = 6). Skin filled mostly with large, oval or triangular plates (L = 228 + 41), rarely star-shaped tables; supporting tables (L = 143 + 11) with moder- ate spire (H = 23 + 11), ratio of L to TH = 2.1 + 0.4; podia endplates (130-160). Usually small (L = 111 + 18) circu- lar tables with small two-pillared spire; supporting tables (L = 96 11) with medium spire (H = 19 5) often topped with 2 “‘horns’’; ra- tio of L to TH = 2.1 + 0.4; podia endplates (114—123, n = 3). Thick oval, knobbed, perforated plates (L = 147 + 34, n = 20) with meshwork of bumps covering one side; curved supporting tables with wide low spire covering half length; well developed end-plates (120-140, n = 4). Crowded layer of acornlike cups (~60) with 2 pillared, tapering spire rising from a cupshaped base; curved supporting tables with medi- um tall spire topped with few blunt teeth; podia with large endplates. lhe [AP Circular to triangular or star-shaped tables (L = 194 + 45, n = 140) with a wide central spire (width at base 102 + 32, n = 140); support- ing tables (L = 110 + 15, n = 90) with low bumpy spire (H = 16 + 5, n = 90), ratio of L to TH 1.9 + 0.3; podia endplates (115-135, n = 5). Introvert Elongate diamond- shaped plates with 2 larger central holes, some with bumps or low pillar arch Oval perforated plates with serrated edge and blunt spines on surface Delicate perforated plates with a few knobs; approaching rosettes Oblong reticulated plates (Bergen) Unknown Elongate oval perfora- ted plates with nu- merous bumps and in some, raised cen- tral rods; two larger central holes 545 Tentacles Ten (8:2); ossicles long thin curved rods and narrow perforated plates Ten; 8:2 small, 1 spec had 9:1 small. Dark around mouth, brown specks on oral disk and tenta- cles 8:2 small; brown spots; oblong plates and rods (Bergen) 8:2 tentacles; various sized plates often with two enlarged central holes Tentacles missing 8:2 small; speckled brown; ossicles of two forms, finer curved, elongate oval plates and large ro- bust rods with a few holes 546 Table 1.—Continued. Pentamera species Distribution P. pediparva, new species Fitz Hugh Sound, B.C. to northern California; 7-120 m brown in preserved state. Podia in five se- ries, of up to 8 rows each at the midbody. Ten small dendritic tentacles including smaller ventral pair. Calcareous ring long and tubular with forked tails on the radials; a mosaic of smaller segments; ratio of in- terradials to radials approximately 1:3. Skin ossicles small oval tables (<100 wm di- ameter) with a simple, two-pillared spire in the form of a low arch; 4 central holes ad- join the spire. Curved supporting tables (length about 150 pm) with low spire. In- trovert has smaller oval tables with spires having a spiny tip. Tentacle ossicles in two forms, finer, curved oval plates and large robust rods with a few larger holes. End plates of podia 200—250 wm in diameter. Material examined.—Twenty-five speci- mens from 12 localities in British Colum- bia, two localities in Washington and one in California. Number of specimens in pa- rentheses after the catalogue number; depth in metres (m). Holotype.—RBCM 997-124-1 (1), length 8 cm, female, collected by Philip Lambert with SCUBA, station L80-50, 29 June 1980. Type locality.—British Columbia, Van- couver Island, Quatsino Sound, Forward In- let, Hall Bank, 50°29.7'N, 128°1.5'W, 7.5 m, cobble-gravel. Paratypes.—RBCM 980-333-100 (1), length 8.2 cm, male, collected by P. Lam- bert with scuba, station L80-50, 29 Jun 1980, British Columbia, Vancouver Island, Quatsino Sound, Forward Inlet, Hall Bank, 50°29.7'N, 128°1.5'W, 7.5 m, cobble-grav- el. RBCM 981-197-2 (1), length 2.5 cm, male, collected by P. Lambert aboard Strickland, Station L81-13, 20 Aug 1981, Curved and tapering at ends; skin wrin- kled; not as stiff as P. rigida PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Body shape Podia Colour Numerous fine Whitish podia in 5 bands, re- duced at oral and anal ends British Columbia, Fitz Hugh Sound, Fair- mile Pass, 51°38'N, 127°51 We) 70mm CMNI 1998-0001 (1), length 7.4 cm, fe- male, collected by D. B. Quayle, 14 Jul 1946, British Columbia, Klucksiwi, Sey- mour Inlet, 51°05’N, 127°39’W, 82 m, sand- gravel. RBCM 973-235-24 (1), length 3.5 cm, sex undet., collected by P. Lambert with scuba, 14 Aug 1973, British Columbia, Barkley Sound, Treble Island, 48°56'N, 125°16.9'W, 9-12 m, sand-shell slope. RBCM 987-253-11 (1), length 3.2 cm, col- lected by G. Green and K. Sendall on Par- izeau, Station G87-17, 23 Jul 1987, Wash- ington, off Cape Flattery, 48°20.9’N, 125°21.39'W, 115-120 m, gravel. CASIZ 50222 (2), collected by Lloyd Barker, 13 Nov 1970, California, Eureka, west of Big Lagoon, 41°10.7'N, 124°7.0'W, 18-37 m. Other collections.—British Columbia: RBCM 982-326-2 (1), on G. B. Reed, FRB station 70-24, Jul 1970, Bramham Island, 51°05'N, 127°39'W, 82 m, sand-gravel. RBCM 977-444-8 (4), collected by D. B. Quayle, FRB station 63-2, 12 Jan 1963, Cormorant Channel, Malcolm Island, 50°36.9’N, 126°57.7'W, 26 m, sand. NMC 1979-1529 (1), FRB St. 18, Comox Bay 49°39'N, 124°55'W, 27 m. RBCM 975-189- 5 (1), formerly BMS 496 (1), St.91/75, 18 Apr 1975, Barkley Sound; Verbeke Reef, 48°52.2'’N, 125°22.6’W, 31—46 m, cobble and gravel. RBCM 984-246-1 (6), FRB #8, 23 Jun 1961, Vancouver Island, Barkley Sound, 48°49'N, 125°34.3’W, 73 m, mud- gravel. RBCM 106-49 (1), collected by D. Ellis, 11 Mar 1965, Saanich Inlet, Mill Bay, 48°38.8'N, 123°32.3'W, 18 m, sand. RBCM 975-294-5 (2), collected by D. Ellis, 3 Nov VOLUME 111, NUMBER 3 Table 1.—Continued. Extended. 547 Calcareous ring Mid-dorsal Ossicles Introvert Tentacles Very long posterior Small, round tables (L = 70 + 11, n = 70) with 4 main holes and 4 or more smaller holes, low 2-pillared spire; in addition, larger diamond- shaped tables (L = 123 + 21,n = 29): slightly curved supporting ta- bles (L = 139 + 18, n = 100) with processes; 1:3 Tables similar to skin but slightly smaller and with teeth on spire 8:2; 2 types of ossicles: finer oblong, curved, perforated plates with 2 larger central holes; and large ro- bust curved rods with holes low spire (H = 19 + 8, n = 100); ratio of L to TH 2.7 + 0.6, n = 100; podia with large endplates (205-215, n = 3). 1965, Vancouver Island, Saanich Inlet, Mill Bay, 48°38.8'N, 123°32.3'W, 18 m, sand. Washington: 1 specimen, Station WP215N Rep A, collected by Alan Fuku- yama, 7 Oct 1996, 66 m north of West Point outfall, Seattle, Puget Sound, 47°39.6'N, 122°26.8'W, 230 m. 1 specimen, AL172N Rep A, collected by Alan Fukuyama, 7 Oct 1996, 52.4 m north of Alki Point outfall, Seattle, Puget Sound, 47°34.2'N, 122°25.3’'W, 143 m. Description.—Total length 1.3—8.2 cm (measured along the outer curve of the body); mean length 3.3 cm for 16 speci- mens. Holotype 8 cm long (Fig. 4). Body curved with a blunt anterior end and taper- ing posterior end. Skin of interambulacral area soft with transverse wrinkles. Five bands of podia each with 4 irregular crowd- ed rows more sparse distally. Podia cylin- drical, non-retractile and bristling with os- sicles. In alcohol, small specimens white with yellowish podia; larger specimens off- white. Ten dendritic tentacles including small ventral pair. Brown spots near base of ten- tacles and in entrance to mouth. Madreporite in shape of a flattened hemi- sphere lies in the dorsal mesentery usually two-thirds of the distance from the anterior end of the calcareous ring. Narrow, con- voluted stone canal attaches to the circular water ring at the posterior tips of the cal- careous ring. One polian vesicle, usually on the ventral side. Two respiratory trees emerge from cloaca and immediately split into two branches, the dorsal branch longer than the ventral. Thin-walled cloaca about one-third of body length and attached to the body wall by numerous strands or muscles. Retractor muscles fairly stout compared to P. rigida; attached to longitudinal mus- cles about % of the distance from anterior end of body. Two tufts of unbranched gonadal tubules attach to dorsal body wall about one-third of the way from anterior end of body, one tuft on each side of the dorsal mesentery. Calcareous ring takes up a large propor- tion of internal body cavity and in severely contracted specimens may appear to span the length of the cavity. Posterior tails of radials long and thin; may be bent back on themselves at posterior end. Transverse thickenings indicate joints between mosaic of pieces. Ratio of interradial to radial length about 1:3 (Fig. 5F). Ossicles of mid-skin small oval or squar- ish tables (diameter 70 pm + 11 SD, n = 71) with a low spire consisting of two pil- lars (Fig. 5A). Pillars may be joined at the top and have one or two side branches, or appear as just two bumps. Most tables have four main holes around the spire with one to several secondary holes. Among oval ta- bles are diamond-shaped tables (length 123 + 21, n = 30) with a spire shaped like a low arch. Toward anterior and posterior ends and in the introvert, spires of tables 548 have multiple spines at apex and tables slightly smaller on average (Fig. 5B). Sup- porting tables of podia curved rods (length 139 + 18, nm = 100) with a low central spire (height of spire 19 wm + 8, n = 100) with two or three bumps, in side view (Fig. 5E). Podia have large end-plates (diameter 205— 215, n = 3) (Fig. 5C). Five anal teeth sur- round anus. Tentacles have two types of ossicles: large, elongate, curved perforated rods and more delicate elongate-oval or curved plates perforated with smaller holes and serrated around edges (Fig. 5D). Etymology.—The species name, pedipar- va, is based on the Latin words pedis mean- ing foot and parvus meaning small or little, referring to the numerous, small podia. Distribution and habitat.—P. pediparva is known from Fitz Hugh Sound, on the central British Columbia coast (51°38’N, 127°51’W), south to Eureka, California (41°10.7'N, 124°7.0’W) (Fig. 6). It ranges in depth from 8 to 120 m on the continental shelf. Usually dredged from firm substrata such as mud-gravel and sand-gravel. Discussion Table 1 compares the features of 14 spe- cies of Pentamera from the west coasts of North and South America, including the two new species described in this paper. New information, not included in original descriptions, has been added to complete the table. Pentamera rigida is allied to P. populi- fera in the shape of the table ossicles, but those of P. rigida have a broad low spire and a more triangular rather than star- shaped plate. P. rigida seems to prefer a firm sand or gravel substratum while P. po- pulifera is usually more common in mud. P. rigida also resembles P. pseudopopuli- fera in general appearance, but the tables are about one-half the diameter of P. rigida and the central spire is much smaller and more delicate. In general appearance, Pentamera pedi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON parva resembles P. lissoplaca with soft skin and fine podia in five rows, but the skin ossicles of the two species differ. P. lisso- placa has a few, tiny tables scattered among the dominant diamond-shaped plates with- out spires. P. pediparva has approximately equal proportions of oval tables and dia- mond-shaped tables of similar size. The curved supporting tables of P. lissoplaca have a moderately tall spire while those of P. pediparva have a low spire. Pentamera constricta (Ohshima 1915) from Japan has tables similar to P. pediparva but it also possesses large smooth plates and the skin is stiff and rough to the touch. The known distributions of P. rigida and P. pediparva are within the Oregonian Province with a southern boundary near Point Conception, California. The northern boundary of this region is poorly defined and varies with the taxa being considered (see Lambert 1996) for discussion. The cluster of specimens in the British Colum- bia region is probably more indicative of collecting pressure than true abundance. I suspect that these two species may be mixed in with specimens of the three close- ly related species P. populifera, P. pseu- dopopulifera and P. lissoplaca in other mu- seum collections. Also, a lack of collecting along the outer coasts of Washington and Oregon could explain the distribution gap for P. pediparva. The two new species are included in the following key to the 14 species of Penta- mera of the west coast of North and South America. This key is based primarily on os- sicles. See Fig. 7 for explanation of dimen- sions of supporting tables used in the key. All measurements in pm. Key to the Pentamera of the West Coast of North and South America 1(0) Mostly round to oval plates without spires, in skin; table ossicles rare or absent. — Mostly large angular or star-shaped plates without a spire; occasional ta- bles Pentamera pseudocalcigera VOLUME 111, NUMBER 3 Fig. 7. 549 L, length. 2(1) Mostly diamond-shaped plates; a few minute tables .... Pentamera lissoplaca Only table ossicles with spires in skin Small oval plates with 2 central and up to 8 marginal knobs. Length of plates between 0 and 99 pm ....... Re Pinay og exc vs cdl aet Pentamera montereyensis Thick, oval, knobbed, perforated plates; mean length 100 to 149 wm; diameter of podial end-plates 100 to 1G Ao alee eRe et eae hee Pentamera trachyplaca Mostly oval plates; mean length be- tween 150 and 199; diameter of podial - end-platesr1>0"to 199M. 2 tee eee . RNS ine ore ee See Pentamera calcigera Supporting tables present in podia .. 4 Supporting tables absent .......... Font eee eee Pentamera chierchia PUEWUG OO Ciorhererie 0 oivks cee Cae oem eo a 5 Spire medium high, between 25 and SER Ae PRA SC ESOS HELE ALA Pda ie dns FETS BAS 6 Spire tall, between 50 and 74 ...... 8 Supporting tables with L: TH ratio of 1.5 to 1.9; mean length of tables 150 to 199; introvert with oval plates Ale FoAtay & ct SAE. AES SAS os Pentamera rigida Supporting tables with L: TH ratio of 2 to 2.4; mean length of tables 100 to 149; introvert with rosettes ........ SRO best he 3 Pentamera pseudopopulifera Supporting tables with L: TH ratio of Abbreviations for dimensions of supporting tables: TH, total height; H, height; C, height of curve; 2.5 to 2.9; mean length of tables 50 to 99; introvert with tables ........... aA Rh AD ee EL Pentamera pediparva 6(4) Supporting tables with L: TH ratio of tO el Ae Oe ON Serene oe. Cesenet i — Supporting tables with L:TH ratio of HES Ow ORNs ce omen Pentamera chiloensis 7(6) Mean length of supporting tables 50 to 99; body U-shaped .... Pentamera zacae — Mean length of supporting tables 100 to) l49-;body curved) sian a sts - seats I ey ae aire atic ie Pentamera populifera 8(4) Mean length of tables less than 50; width of spire at base less than 25 .. ci Siecle Nom AEG Pentamera beebei — Mean length of tables 50 to 99; Width of spire at base 25 to 49 .......... Aiea, JN earn as Pentamera charlottae Acknowledgments I am grateful to G. Green and K. Sendall of the Royal B. C. Museum for the field assistance. Many specimens in the RBCM were collected by the late Dan Quayle and Frank Bernard of the Pacific Biological Sta- tion, who diligently collected marine inver- tebrates in the 1950’s and 60’s. I would like to pay tribute to them for their foresight and contributions to our knowledge of inverte- brates in British Columbia. Thanks also to 550 G. Hendler and K. Groves, LACM and B. van Syoc and E. Kools, CASIZ, for loans of specimens. A. Fukuyama of the Univer- sity of Washington kindly loaned me two lots of Pentamera from one of his surveys. Thank you to Dr. D. Pawson and an anon- ymous reviewer for their constructive com- ments. Support from the Executive Director of the Royal British Columbia Museum for this taxonomic research is gratefully ac- knowledged. Literature Cited Ayres, W. O. 1852. Description of a new species of Holothuria.—Proceedings of the Boston Society of Natural History 4:207—208. Bergen, M. 1996. 9. Class Holothuroidea: Including keys and descriptions to all continental shelf species from California. Pp. 195-250 in J. A. Blake, P. H. Scott, & A. Lissner, eds., Taxonom- ic atlas of the benthic fauna of the Santa Maria Basin and the Western Santa Barbara Channel, Santa Barbara Museum of Natural History, Santa Barbara, 305 pp. Clark, H. L. 1924. Some holothurians from British Co- lumbia.—Canadian Field Naturalist 38:54—57. Deichmann, E. 1938a. Eastern Pacific expeditions of the New York Zoological Society. XVI. Holo- thurians from the western coasts of Lower Cal- ifornia and Central America, and from the Ga- lapagos Islands.—Zoologica (New York) 23: 361-387. . 1938b. New holothurians from the western coast of North America and some remarks on the genus Caudina.—Proceedings of the New England Zoological Club 16:103-115. . 1941. The holothurioidea collected by the Ve- lero II during the years 1932 to 1938. Part 1, Dendrochirota.—Allan Hancock Pacific Expe- ditions 8:61—195. Lambert, P. 1985. Geographic variation of calcareous ossicles and the identification of three species of eastern Pacific sea cucumbers (Echinoderma- ta: Holothuroidea). Pp. 437—443 in B. FE Keegan & B. D. S. O’Connor, eds., Echinodermata: Pro- ceedings of the Fifth International Echinoderm PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Conference, Galway, A. A. Balkema, Rotter- dam, 662 pp. . 1996. Psolidium bidiscum, a new shallow-wa- ter psolid sea cucumber (Echinodermata: Hol- othuroidea) from the northeastern Pacific, pre- viously missidentified as Psolidium bullatum Ohshima.—Canadian Journal of Zoology 74: 20-31. . 1997. Sea cucumbers of British Columbia, Puget Sound and Southeast Alaska. Royal BC Museum and University of BC Press, Vancou- ver, 192 pp. Ludwig, H. 1886a. Echinodermen des Beringsme- eres.—Zoologiscen Jahrbiichern. Zeitschrift fiir Systematik, Geographie und Biologie der Thi- ere 1886:275—296. . 1886b. Die von G. Chierchia auf der Fahrt der Kgl. Ital. Corvette ““Vettor Pisani’ gesam- melten Holothurien.—Zoologiscen Jahrbtich- ern. Zeitschrift fiir Systematik, Geographie und Biologie der Thiere 1886:1—36. Ohshima, H. 1915. Report on the holothurians col- lected by the U.S. fisheries steamer “‘Albatross”’ in the N.W. Pacific during the summer of 1906.—Proceedings of the United States Na- tional Museum 48:213-—291, pls. 8-11. Panning, A. 1949. Versuch einer Neuordnung der Fam- ilie Cucumariidae (Holothurioidea, Dendrochi- rota).—Zoologische Jahrbiicher Abteilung fur Systematik, Oekologie und Geographie der Ti- ere 78:404—470. Pawson, D. 1982. Holothuroidea. Pp. 813-818 in S. P. Parker, ed., Synopses and classification of living organisms, McGraw-Hill, Toronto, 1232 pp. , & H. B. Fell. 1965. A revised classification of the Dendrochirote Holothurians.—Breviora 214:1-7. Stimpson, W. 1851. Descriptions of new species of holothurians.—Proceedings of the Boston So- ciety of Natural History 4:67. . 1864. Descriptions of new species of marine invertebrata from Puget Sound, collected by the Naturalists of the North-west Boundary Com- mission, A. H. Campbell, Esq., Commission- er.—Proceedings of the Academy of Natural Sciences of Philadelphia 16:153—161. Thandar, A. S. 1990. The phyllophorid holothurians of southern Africa with the erection of a new ge- nus.—South African Journal of Zoology 25: 207-223. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):551—603. 1998. The Anacroneuria of Costa Rica and Panama (Insecta: Plecoptera: Perlidae) Bill P. Stark Department of Biology, Mississippi College, Clinton, Mississippi 39058, U.S.A. Abstract.—Twenty seven Anacroneuria species are recorded for Costa Rica and Panama including 18 described as new species (Anacroneuria alajuela, A. benedettoi, A. curiosa, A. exquisita, A. hacha, A. harperi, A. holzenthali, A. marca, A. marginata, A. maritza, A. perplexa, A. talamanca, A. tornada, A. uatsi, A. varilla, A. ventana, A. zapata, A. zarpa). Redescriptions are given for nine species previously recorded from the area. Lectotypes are designated for Anacroneuria expansa Klapalek, A. annulipalpis Klapalek, A. planicollis Kla- palek and A. plutonis (Banks); A. expansa Klapalek = A. acutipennis Klapdlek, A. chiapasa Jewett = A. planicollis Klapalek, A. dampfi Jewett = A. planicollis Klapalek, and A. tristani (Navas) = A. plutonis (Banks) are placed in synon- ymy. A provisional key for males is provided. The stonefly fauna of lower Mesoamerica includes only the speciose neotropical per- lid genus Anacroneuria. Despite sporadic attempts (e.g., Needham & Broughton 1927, Jewett 1958, Harper 1992) this fauna remains virtually undocumented. Eleven of the twelve previously proposed Costa Rican and Panamanian species are known from obscure, scattered and undiagnostic original descriptions published early in this century. At that time the importance of internal male genitalic characters was unrecognized and similarities in size, coloration and subgen- ital plate structure led to the erroneous con- cept of Anacroneuria species with ranges extending from Mexico to Brazil (Needham & Broughton 1927). More recent studies (Zwick 1972, 1973; Stark 1995) suggest the male aedeagal structure is the most reliable structure for species recognition in this ge- nus. Egg shape and size are useful in dis- tinguishing some species but the chorionic surface lacks the detail found in other per- lids. In addition to the 12 species whose ho- lotypes or lectotypes are based on Costa Ri- can or Panamanian material, Needham & Broughton (1927), Jewett (1958) and Harp- er (1992) include records of three earlier described species and records of three spe- cies based on paratypes. Needham & Broughton (1927) and Jewett (1958), for example, both record A. cincta (Pictet) and A. nigrocincta (Pictet) from the region but Zwick (1972, 1973) has shown neither of these can be considered valid records. Be- cause the holotype of A. cincta is lost, that species is considered a nomen dubium, and the species identified as A. nigrocincta in these studies applies to another, presently unidentified species. Harper’s (1992) record of A. cincta from Panama is based on Need- ham & Broughton’s (1927) invalid defini- tion of the species. Jewett (1958) included Panamanian specimens among the para- types of A. crenulata Jewett and A. flavom- inuta Jewett. Harper’s (1992) figures of the male aedeagi of these specimens indicate they are distinct from the respective allo- types. These specimens are assigned to two of the new species described in this study. Unfortunately the specimen listed by Harp- er (1992) as the putative male of A. pallida Jewett could not be located. Anacroneuria sulana Needham & Broughton (1927), re- corded by Jewett (1958) from Panama, was S52 placed as a synonym of A. annulicauda (Pictet) by Zwick (1972). Jewett’s Pana- Manian specimens, presumably in the American Museum of Natural History, need to be reexamined before this identification is accepted. During the late 1980’s and early 1990’s, R. W. Holzenthal and colleagues at the Uni- versity of Minnesota and D. H. Funk, J. Jackson and colleagues at the Stroud Water Research Center made extensive light trap collections of Costa Rican stoneflies. This valuable material was made available for my study and has been supplemented with specimens from the museums listed below. Results from available material indicate a minimum of 27 species occur in Costa Rica and Panama. This includes nine of the ear- lier named species and 18 that are new to science. A relatively strong pattern of en- demism emerges from this study with 21 species known only from Costa Rica-Pan- ama. None of the species reported in Costa Rica or Panama are known to occur in South America and only two are currently reported from Mexico (Jewett 1958, Stark 1995). Males of thirteen species were taken in light traps from a single locality at Es- tacion Maritza in Guanacaste Province. Materials and Methods Aedeagai and female terminalia were prepared by clipping the abdomen and plac- ing it in a small beaker of KOH. The so- lution was brought to a boil, then the ter- minalia were placed in dishes of distilled water and extraneous tissue removed with fine forceps. The aedeagus was also re- moved from the abdominal cavity with fine forceps and figures were prepared in dorsal, ventral and lateral aspects at 100; figures of the male ninth sternum were drawn at 50X and the female terminalia at 25x. Fig- ures of the adult head and pronota were drawn at 25X from specimens in alcohol. Aedeagal preparations for scanning elec- tron microscopy were made by crudely dis- secting the structure from specimens in al- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cohol. The aedeagi were sonicated in ace- tone for one minute, air dried, placed on specimen stubs with double stick copper tape, sputter coated with gold-palladium, and the apex examined in ventral and lateral aspect with an AMRAY 1810D scanning electron microscope. Holotype specimens of new species are placed in the United States National Mu- seum (USNM); some specimens designated as holotypes are placed in the United States National Museum through the courtesy of Brigham Young University, Stroud Water Research Center, University of Minnesota and Utah State University. Paratypes and other specimens are deposited in the fol- lowing collections as indicated: Bill P. Stark, Clinton, Mississippi (BPS); Brigham Young University, Provo, Utah (BYU); C. P. Gillette Museum of Arthropod Diversity, Colorado State University, Fort Collins, Colorado (CSU); Cornell Univer- sity, Ithaca, New York (CU); Florida State Collection of Arthropods, Gainesville, Flor- ida (FSCA); Instituto Nacional de Biodiv- ersidad, Santo Domingo, Costa Rica (IN- BIO); Museum National d’Histoire Natu- relle, Paris (MNHN); Museum of Compar- ative Zoology, Harvard University (MCZ); National Museum of Natural History, Prague (NMP); Ohio State University, Co- lumbus, Ohio (OSU); Stroud Water Re- search Center, Avondale, Pennsylvania (SWRC); University of Minnesota, Saint Paul, Minnesota (UMSP); Utah State Uni- versity, Logan, Utah (USU). Provisional Key to Regional Male Anacroneuria 1. Median pronotal band brown (Fig. 36) ots Buslbiantie -oysean eat tone eee enn Z — Median pronotal band yellow, or pro- notum without distinct pigment bands Siete a aisha, tin Shag ieee ee eae 7 2. Hammer a low indistinct mound (Fig. AR cee eRe eee eee hacha — Hammer well developed, thimble or nipple*shaped '(Figs. 7962) sop. 3 3. Aedeagal apex projecting ventrally be- VOLUME 111, NUMBER 3 TS Nn Ne) 10 11 12 tween hooks (Fig. 108); dorsal aedeagal keel absent tornada Aedeagal apex not projecting between hooks (Fig. 93); dorsal aedeagal keel well developed (Fig. 94) ........... 4 Aedeagal apex with three subequal LODESR EIS, QD) ie. ee ce soe planicollis Aedeagal apex simple or with minute lateral lobes (Figs. 65, 85) ......... 5 Aedeagal apex beyond shoulders at least twice as long as wide (Fig. 84) 5 3 bib .ot S15 Onc G Smee ein eer te Saar eas maritza Aedeagal apex beyond shoulders about as long as wide (Fig. 64) .......... 6 Lateral aspect of aedeagal apex wider than long at widest point (Fig. 38) divisa Lateral aspect of aedeagal apex longer famewiGes(E18: OS). 6 qe sa lineata Hammer poorly developed or absent ORRA Sts cast: Sonica Seek es 8 Hammer thimble shaped, well devel- OPC OMCs) eames ohne oes als te shee 12 Forewing length 7-11 mm ......... 9 Forewing length at least 14 mm ..... 10 Aedeagal apex trilobed, lateral lobes small, hooks chelate (Fig. 90) ... perplexa Aedeagal apex simple, hooks slender (Fig. 130) varilla in part Hammer absent (Fig. 67); aedeagal apex simple, usually with a sharp trans- verse keel (Fig. 68) magnirufa Hammer a low obscure mound (Fig. 2); aedeagal apex trilobed, without keel (EVE, 48) Gre AA a ene A RATA Sena 11 Mesal lobe of aedeagal apex deeply notched (Fig. 4); wing membrane with- out transparent circular spot near cord acutipennis Mesal lobe of aedeagal apex rounded (Fig. 44); wing membrane with trans- parent circular spot near cord .. exquisita Head without brown pigment spot be- tween or just anterior to ocelli (Fig. 11) — Head with brown pigment between or 13) just anterior to ocelli (Figs. 76,96) .. 16 Aedeagal apex and hooks asymmetrical (BiSSESB=55D) Mie ett. eee atets harperi — Aedeagal apex and hooks symmetrical 14. Median yellow pronotal stripe less than a fourth of pronotal width (Fig. 20. De Dp 13 393) 131); aedeagal apex with clawlike dor- solateral lobes (Fig. 134) zarpa Median yellow pronotal stripe about half of pronotal width (Figs. 11, 31); aedeagal apex without clawlike dorso- lateralWlobes nic, 2528). Ke PROS we. 15 Dorsal aedeagal apex acute and simple (Fig. 14) annulipalpis Dorsal aedeagal apex rounded and tri- lobedi(Fige S34) Mii. 22 aes curiosa Aedeagal apex massive with winglike shoulders (Figs. 20, 60) Aedeagal apex small and without winglike shoulders (Fig. 114) ....... 18 Lateral aedeagal wings large and pointed (Fig. 60) holzenthali Lateral aedeagal wings small and rounded (Fig. 20) benedettoi Forewing length at least 13 mm .... 19 Forewing length no more than 10 mm Aedeagal apex with small dorsolateral lobes, apex shoe shaped in lateral as- pect (Figs. 128-129) zapata Aedeagal apex without dorsolateral lobes, lateral aspect not shoe shaped Dorsal aedeagal keel well developed (Fig. 124); wing membrane usually with a circular transparent spot at cord Sed ates. oes aia cee cet ca ventana Dorsal aedeagal keel absent (Fig. 74); wing membrane without circular trans- PAKENGSPOl Gites 5 ee tw eet, aecle 21 Aedeagal apex gradually narrowed to tip, bases of aedeagal hooks bulging (Fig. 100); M-line and lateral callosi- ties on head distinct (Fig. 96) .. plutonis Aedeagal apex with almost parallel sides distal to shoulders, bases of hooks not enlarged (Fig. 75); head pat- tern with M-line and callosities indis- tinct (Fig. 71) marca Ventral aedeagal apex with prominent pair of membranous lobes (Fig. 120) Ventral aedeagal apex without mem- branous lobes (Fig. 10) Aedeagal apex offset from shoulders by lateral notches, tip acute (Fig. 24) SAE ag SNe LU es, «MN Stee Soe blanda Aedeagal apex without lateral notches, 554 tip truncate or emarginate (Fig. 119) of. . gE SEY eee varilla in part 24. Aedeagal apex projecting fingerlike from shoulders (Figs. 114-115) .... uatsi — Aedeagal apex not fingerlike, lateral margins more or less convergent to tip (Figs. 10, 80) 25. Aedeagal apex with a U-shaped dorsal keel (Fig. 9) alajuela — Aedeagal apex without U-shaped keel 26. Aedeagal apex gradually narrowed from bases of hooks to near tip (Fig. MA!) pt Sen ANE cores. iene x vgs talamanca — Aedeagal apex not conspicuously nar- rowed beyond hooks (Figs. 29, 80) .. 27 27. Aedeagal apex with well developed dorsal keel (Fig. 79); area beyond hooks about as wide as long (Fig. 80) eS ee Doo ana marginata — Aedeagal apex with obscure dorsal keel (Fig. 29); area beyond hooks lon- gen than wide (Eis. 30) fess. - costana Anacroneuria acutipennis Klapalek Figs. 1-5, 136, 159-160 Anacroneuria acutipennis Klapalek, 1923: 23. Holotype 2, Volcan Chiriqui, Pana- ma (NMP). Anacroneuria expansa Klapalek, 1923:22. Lectotype 2, Guatemala (NMP), new synonymy. Anacroneuria sp. C: Harper, 1992:118. Adult habitus.—Head yellow except for lappets and dark area forward of ocelli. Pale median pronotal stripe wide, midlateral dark stripes irregular (Fig. 1). Wing mem- brane transparent, veins pale brown; C and Sc pale. Male.—Forewing length 16-17 mm. Hammer a low mound (Fig. 2). Aedeagal apex trilobed, mesal lobe deeply notched. Hooks stout, dorsal keel absent (Figs. 3—5, 159-160). Female.—Forewing length 19-21 mm. Subgenital plate bilobed, lobes broad and usually emarginate; notch V-shaped. Trans- verse sclerite of sternum nine well devel- oped; mesal sclerite T-shaped, lateral setae prominent (Fig. 136). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Nymph.—Unknown. Material.—Costa Rica: Alajuela: Rio Peje, 1 km SE San Vicente, 1450 m, 14 Feb 1992, R. Holzenthal, EK Munoz, K. Kjer, 1 3, 1 2 (UMSP). Cartago: Reserva Tapanti, Quebrada Palmitos, 1400 m, 2 Jun 1990, R. Holzenthal, R. Blahnik, EF Munoz, 1 6, 3 2 (BPS). Same location, 24 Mar 1991, R. Holzenthal, EF Munoz, J. Huisman, 3 6 (IN- BIO). Same location, 1 Aug 1990, 3 @ (UMSP). Puntarenas: Rio Bellavista, ca. 1.5 mi NW Las Alturas, 1400 m, 8 Apr 1987, R. Holzenthal, S. Hamilton, M. Heyn, 1 6 (USNM). San Jose: Parque Nacional Brau- lio Carrillo, Rio Zurqui, 1650 m, 6 Jul 1986, R. Holzenthal, M. Heyn, B. Armi- tage, 1 d (BPS). Guatemala: No locality data, A. expansa lectotype 2 (NMP). Pan- ama: Volcan Chiriqui, 1907, V. D. Whede, holotype 2 (NMP). Same locality, 5500’, 11 Jan 1964, S. L. Wood, 2 6 (BYU). Comments.—A female specimen bearing label data “‘Anacroneuria expansa Klap. 1923, guat/de Selys 21” from the Natural History Museum, Prague, is selected as lec- totype. Although the forewing length of this specimen is given as 28 mm (Klapdalek 1923) it is similar in body size to other specimens of A. acutipennis, and it is indis- tinguishable from these specimens in sub- genital plate shape and features of sternum nine and thus is placed as a synonym. The Panamanian material recorded as “‘Anacro- neuria sp. C”’ by Harper (1992) is also as- signed to this species. Anacroneuria alajuela, new species Figs. 6-10 Adult habitus.—Head with a large dark area over ocelli extending to M-line; lap- pets brown. Median pronotal stripe yellow, lateral stripes brown (Fig. 6). Wing mem- brane brown, veins brown. Male.—Forewing length 9 mm. Hammer thimble shaped, height greater than basal diameter (Fig. 7). Aedeagal apex broad, scoop shaped and bearing a dorsal U- VOLUME 111, NUMBER 3 S55 Figs. 1-5. A. acutipennis structures. 1. Head and pronotum. 2. Male sternum 9. 3. Aedeagus, lateral. 4. Aedeagus, dorsal. 5. Aedeagus, ventral. Scales: 0.6 mm (1), 0.3 mm (2), 0.15 mm (3-5). shaped keel; hooks moderately thickened Etymology.—The species name, based on (Figs. 8-10). the Costa Rican province in which the ho- Female.—Unknown. lotype was collected, is used as a noun in Nymph.—Unknown. apposition. 556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 6-10. A. alajuela structures. 6. Head and pronotum. 7. Male sternum 9. 8. Aedeagus, lateral. 9. Ae- deagus, dorsal. 10. Aedeagus, ventral. Scales: 0.6 mm (6), 0.3 mm (7), 0.15 mm (8-10). VOLUME 111, NUMBER 3 Types.—Holotype ¢ from Costa Rica, Alajuela, 20 km S Upala, June 1990, E D. Parker (USNM). Diagnosis.—The broad aedeagal apex is similar to that of A. divisa (Figs. 38—40) and A. lineata (Figs. 63—65) but these spe- cies have dark mesal pronotal stripes, low moundlike hammers and narrow longitudi- nal aedeagal keels. Anacroneuria annulipalpis Klapalek Figs. 11-15, 137, 161-162 Anacroneuria annulipalpis Klapalek, 1922: 91. Lectotype ¢, Chiriqui, Panama (NMP). Anacroneuria quadriloba Jewett: Harper, 1992-107. Adult habitus.—Head yellow. Pronotum yellow except for irregular, dark lateral Stripes (Fig. 11). Wing membrane pale am- ber, veins C, Sc pale, others brown. Male.—Forewing length 12-16 mm. Hammer thimble shaped (Fig. 12). Aedea- gal apex greatly narrowed into curved Spine; ventral membranous processes pres- ent. Dorsal keel absent, hooks scooped at apex (Figs. 13-15, 161-162). Female.—Forewing length 16-20 mm. Subgenital plate four lobed; outer lobes broad, but subequal in length to inner lobes. Transverse sclerite of sternum nine curved, mesal sclerite T-shaped and sparsely setose (Fig. 137). Nymph.—Unknown. Material.—Costa Rica: Alajuela: Reser- va Forestal San Ramon, Rio San Lorencito, 980 m, 30 Mar 1987, R. Holzenthal, S. Hamilton, M. Heyn, 1 6 (UMSP). Same location, 1 May 1990, R. Holzenthal, R. Blahnik, 1 2 (UMSP). Guanacaste: Esta- cion Maritza, Rio Tempisquito, 5 May 1989, 7 3 (BPS). Same location, 11 May 1990, 1 6, 1 2 (SWRC). Same location, 30 May 1990, 2 ¢ (SWRC). Same location, 22 Oct 1990, 5 d (INBIO). Puntarenas: Zona Protectora Las Tablas, Rio Coton, Sitio Co- ton, 1460 m, 15 Apr 1989, R. Holzenthal, R. Blahnik, 1 ¢, 1 2 (INBIO). Rio Bella- 557 vista, ca. 1.5 km NW Las Alturas, 1400 m, 2 Aug 1987, R. Holzenthal, J. Morse, P. Clausen, 1 6, 3 2 (UMSP). Rio Jaba, 1.4 km W Las Cruces, 1150 m, 14 Jun 1986, R. Holzenthal, M. Heyn, B. Armitage, 1 3 (BPS). San Vito de Java, 23 May 1960, C. FE Dowling, 1 6 (FSCA). San Jose: Parque Nacional Braulio Carrillo, Rio Zurqui, 1650 m, 6 Jul 1986, R. Holzenthal, M. Heyn, B. Armitage, 2 ¢, 1 2 (UMSP). El Salvaje, Rio Tabarcia, 8 km E Palmichal, 1650 m, 19 Jan 1992, R. Holzenthal, K. Kjer, FE Quesada, 2 ¢ (INBIO). Panama: Chiriqui Province, 5 km NE El Hato del Volcan, 1 Oct 1975, D. Chandler, 1 6 (BYU). Volcan Chiriqui, 5500’ 11 Jan 1964, S. L. Wood, 2 6 (BYU). Chiriqui, 2 2 syntypes (NMP). Comments.—Two female syntypes were located, each with damaged subgenital plates. One specimen is so severely dam- aged as to prevent diagnosis. The other has the left half of sternum eight and most of sternum nine intact and is chosen as lecto- type. Both specimens are in the National Museum of Natural History, Prague and were studied while on loan to P. Zwick. Fe- male A. quadriloba Jewett are indistin- guishable from this species but the aede- agus of paratopotypes are more similar to those of A. lineata (Fig. 95). However, the Panamanian records listed for A. quadrilo- ba by Harper (1992) are to be included with this species. Costa Rican specimens show somewhat more size variation than has been noted for other Anacroneuria. Anacroneuria benedettoi, new species Figs. 16—20, 138, 163-164 Adult habitus.—Head mostly yellow brown with diffuse brown over ocelli and lappets. Median yellow pronotal stripe bor- dered by irregular, broad midlateral brown bands; anterolateral margins pale (Fig. 16). Wing membrane brown, veins brown. Male.—Forewing length 10 mm. Ham- mer thimble shaped, height subequal to di- ameter (Fig. 17). Aedeagal apex trilobed, 558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ~p3 Sd Tate bale Figs. 11-15. A. annulipalpis structures. 11. Head and pronotum. 12. Male sternum 9. 13. Aedeagus, lateral. 14. Aedeagus, dorsal. 15. Aedeagus, ventral. Scales: 0.6 mm (11), 0.3 mm (12), 0.15 mm (13-15). shoulders projecting laterally. Dorsal keel longer than mesal lobes, mesal notch V- on mesal lobe, hooks slender (Figs. 18-20, shaped. Transverse sclerite of sternum nine 163-164). sinuate; T-shaped mesal sclerite setose, lat- Female.—Forewing length 12-13 mm. eral setae prominent (Fig. 138). Subgenital plate four lobed. Lateral lobes Nymph.—Unknown. VOLUME 111, NUMBER 3 559 18 “19 20 Figs. 16-20. A. benedettoi structures. 16. Head and pronotum. 17. Male sternum 9. 18. Aedeagus, lateral. 19. Aedeagus, dorsal. 20. Aedeagus, ventral. Scales: 0.6 mm (16), 0.3 mm (17), 0.15 mm (18-20). 560 Etymology.—The patronym honors L. A. Benedetto for his generosity in sharing in- formation on Anacroneuria type material. Types.—Holotype 6d from Costa Rica, Guanacaste, Rio Los Ahogados, 11.3 km ENE Quebrada Grande, 470 m, 7 Mar 1986, R. Holzenthal, W. Fasth (USNM). Paratypes: Costa Rica: Alajuela: Laguna Rio Cuarto, 2.8 km N Rio Cuarto, 400 m, 13 Feb 1992, R. Holzenthal, E Munoz, K. Kjer, 2 ¢6 (UMSP). Guanacaste: Rio Gon- gora, 4 km NE Quebrada Grande, 590 m, 21 Jul 1987, R. Holzenthal, J. Morse, P. Clausen, 6 6 (INBIO, UMSP). Rio Agua- cate, 0.5 km E Aguacate, 590 m, 16 Feb 1992, R. Holzenthal, EK Munoz, K. Kjer, 1 6 (UMSP). Rio Orosi, Estacion Pitilla, 700 m, 22 May 1990, R. Holzenthal, R. Blah- nik, 2 ¢ (BPS). Parque Nacional Guana- caste, El] Hacha, Quebrada Alcornoque, 250 m, 26 Jul 1987, R. Holzenthal, J. Morse, P. Clausen, 9 ¢ (BPS). Estacion Maritza, Rio Tempisquito, 28 May 1989, 1 56 (SWRC). Limon: Rio Banano, 16 km WSW Bomba, 150 m, 26 Mar 1987, R. Holzenthal, S. Hamilton, M. Heyn, 8 d (UMSP). Puntar- enas: Reserva Biologica Carara, Quebrada Bonita, 35 m, 18—20 May 1990, R. Holzen- thal, R. Blahnik, 1 ¢6 (UMSP). 2 mi SW San Vito de Java, 22—24 Jun 1964, R. Woodruff, 2 56 (FSCA). San Jose: Reserva Biologica Carara, Rio Carara, Carara, 280 m, 14 Feb 1991, R. Holzenthal, EF Munoz, J. Huisman, 2 ¢, 1 9 (UMSP). Panama: Bocas del Toro Province: Miramar, 3 Apr 1979, H. Wolda, 1 6 (BYU). Diagnosis.—This species is similar to A. holzenthali but has smaller and apically rounded aedeagal wings rather than long and pointed wings typical of A. holzenthali. Anacroneuria blanda Needham & Broughton Figs. 21-25, 139 Anacroneuria blanda Needham & Brough- ton, 1927:117. Holotype ¢, Barro Colo- rado Island, Panama (CU). Anacroneuria blanda: Harper, 1992:117. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Adult habitus.—Head yellow except for lappets and dark area covering ocelli and extending forward and laterally to M-line. Median pronotal stripe yellow, lateral Stripes brown, anterolateral margins pale (Fig. 21). Male.—Forewing length 9 mm. Hammer cylindrical, height greater than diameter (Fig. 22). Triangular aedeagal apex offset from shoulders by transverse dorsolateral grooves; ventral aspect with a pair of mem- branous lobes, lateral aspect sinuate. Dorsal keel absent, hooks somewhat scoop shaped at the tips (Figs. 23-25). Female.—Forewing length 13-14 mm. Subgenital plate bilobed, margins rounded or truncate, notch V-shaped. Transverse sclerite of sternum nine absent, posterior margin concave; mesal sclerite triangular and sparsely setose (Fig. 139). Nymph.—Unknown. Material.—Costa Rica: Alajuela: 20 km S Upala, 5-10 Mar 1991, EF Parker, 2 6 (BYU, USU). Puntarenas: 2.8 mi E Golfito, 3-4 Jul 1967, O. S. Flint, Ortiz, 1 6 (USNM). Panama: Rio Agua Salud, Pipe- line Road, 8—12 Jul 1967, O. S. Flint, Ortiz, 1 ¢ (USNM). Cerro Campana, 11-14 Jul 1967, O. S. Flint, 2 2 (USNM). Barro Col- orado Island, W. C. Allee, Holotype 2 (CU). Comments.—Anacroneuria blanda was known from the holotype female until Harper (1992) assigned the additional Pan- amanian specimens listed above to this spe- cies. Although these specimens have a larg- er area of dark pigment on the head than the holotype, they are indistinguishable in other features and I concur with this place- ment. Anacroneuria costana (Navas) Figs. 26—30 Neoperla costana Navas, 1924:72. Holo- type 6, Costa Rica (MNHN). Adult habitus.—Head yellow with dif- fuse yellow brown over ocelli; lappets brown. Median pronotal stripe pale, irreg- VOLUME 111, NUMBER 3 561 Figs. 21-25. A. blanda structures. 21. Head and pronotum. 22. Male sternum 9. 23. Aedeagus, lateral. 24. Aedeagus, dorsal. 25. Aedeagus, ventral. Scales: 0.6 mm (21), 0.3 mm (22), 0.15 mm (23-25). 562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bie ete Va a ra} iy ERS . - © ~~ Figs. 26-30. A. costana structures. 26. Head and pronotum Aedeagus, dorsal. 30. Aedeagus, ventral. Scales: 0.6 mm (26), 0.3 mm (27), 0.15 mm (28-30). . 27. Male sternum 9. 28. Aedeagus, lateral. 29. VOLUME 111, NUMBER 3 ular midlateral stripes brown, margins pale (Fig. 26). Wings transparent, veins brown. Male.—Forewing length 8.5 mm. Ham- mer thimble shaped, height greater than di- ameter (Fig. 27). Aedeagal apex simple, scoop shaped; dorsal keel weak, hooks slen- der (Figs. 28-30). Female.—Unknown. Nymph.—Unknown. Material.—Costa Rica: Puntarenas: Rio Guineal, ca. 1 km E Finca Helechales, 840 m, 22 Feb 1986, R. Holzenthal, J. Morse, W. Fasth, 1 6 (BPS). Rio Singri, ca. 2 km S Finca Helechales, 22 Feb 1986, R. Hol- zenthal, J. Morse, W. Fasth, 2 ¢6 (UMSP INBIO). Comments.—Anacroneuria costana is a member of a large and difficult species complex found throughout Central and South America. Among Costa Rican spe- cies, A. costana most closely resembles A. marginata, new species, but that species has a shorter and more robust aedeagal apex. The aedeagus of these Puntarenas males closely resembles the figures of the holotype provided by L. Benedetto (L. Be- nedetto, pers. comm.). Anacroneuria curiosa, new species Figs. 31-35, 165-166 Anacroneuria flavominuta: Harper, 1992: eG: Adult habitus.—Head yellow except for lappets. Pronotum with irregular dark lat- eral bands and a wide mesal yellow band (Fig. 31). Femora yellow with narrow black apical band. Wing membrane and most veins brown; R vein dark brown, C pale. Male.—Forewing length 8-9 mm. Ham- mer thimble shaped, height greater than di- ameter (Fig. 32). Dorsal aspect of aedeagal apex trilobed; mesal lobe longer than lateral lobes and weakly keeled basally (Figs. 33- 35, 165-166). Apex offset from dorsal sclerite by prorninent transverse fold (Fig. 165). Female.—Unknown. Nymph.—Unknown. 563 Etymology.—The species name, meaning inquisitive or curious, alludes to the pecu- liar transverse dorsal fold on the aedeagal apex whose function will require even greater curiosity to discern. Types.—Holotype ¢ and 15 6 paratypes from Costa Rica, Guanacaste, Parque Na- cional Guanacaste, Maritza, Rio Tempis- quito, 550 m, 17 Jun 1988, C. M. Flint, O. S. Flint, R. Holzenthal (USNM). Additional Paratypes: Costa Rica: Alajuela: Rio Boch- inche tributary, Cerro Campana, 6 km NW Dos Rios, 600 m, 22 Jul 1987, R. Holzen- thal, J. Morse, P Clausen, 3 6 (UMSP). Guanacaste: Quebrada Garcia, 470 m, 8 Mar 1986, R. Holzenthal, W. Fasth, 1 6 (BPS). Parque Nacional Guanacaste, Esta- cion Pitilla, Rio Orosi, 700 m, 19 Jun 1988, C. M. Flint, O. S. Flint, R. Holzenthal, 2 6 (USNM). Parque Nacional Guanacaste, Es- tacion Maritza, Rio Tempisquito, 550 m, 30 Aug 1990, J. Huisman, R. Blahnik, FE Que- sada, 1 ¢ (UMSP). Estacion Maritza, Rio Tempisquito, 25 May 1990, 1 ¢6 (SWRC). Same location, 30 May 1990, 1 6 (INBIO). Nicaragua: Pte. Quinama, E Villa Somoza, 29 Jul 1967, O. S. Flint, 1 ¢6 (USNM). Diagnosis.—This species resembles A. uatsi in size and general coloration but the two are quite distinct in aedeagal features. In dorsal aspect A. curiosa has a complete transverse fold (Fig. 34) whereas A. uatsi has only a minute keel (Fig. 114). Comments.—This small species ranges at least from Panama to Nicaragua but may be uncommon. Harper’s (1992) figure of the aedeagus of an A. flavominuta paratype from Panama is typical of this species and distinct from the allotype male of A. fla- vominuta from Mexico. Anacroneuria divisa (Navas) Figs. 36—40, 141, 167-168 Forquilla divisa Navas 1924:74. Syntype 2, Costa Rica (NMNBH). Adult habitus.—Head with dark pattern over ocelli extending onto clypeus, but in- terrupted by pale M-line; pale oval areas 564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 31-35. A. curiosa structures. 31. Head and pronotum. 32. Male sternum 9. 33. Aedeagus, lateral. 34. Aedeagus, dorsal. 35. Aedeagus, ventral. Scales: 0.6 mm (31), 0.3 mm (32), 0.15 mm (33-35). VOLUME 111, NUMBER 3 565 Figs. 36-40. A. divisa structures. 36. Head and pronotum. 37. Male sternum 9. 38. Aedeagus, lateral. 39. Aedeagus, ventral. 40. Aedeagus, dorsal. Scales: 0.6 mm (36), 0.3 mm (37), 0.15 mm (38—40). 566 lateral to ocelli and minute spots anterolat- eral to ocelli, lappets diffuse brown. Me- dian pronotal stripe brown; anterior and posterior margins brown, disc and antero- lateral margins pale (Fig. 36). Wing mem- brane brown, veins C, Sc and R pale, other veins brown. Male.—Forewing length 13-15 mm. Hammer low, nipple shaped (Fig. 37). Ae- deagal apex simple, scoop shaped with broad, rounded shoulders. Lateral aspect expanded through dorsal keel; hooks slen- der (Figs. 38—40, 167-168). Female.—Forewing length 17-19 mm. Subgenital plate four lobed; lateral lobes larger than median lobes. Transverse scler- ite of sternum nine with median notch; me- dian sclerite T-shaped, lateral setae promi- nent (Fig. 141). Nymph.—Unknown. Material.—Costa Rica: Alajuela: Cerro Campana, Rio Bochinche tributary, 6 km NW Dos Rios, 600 m, 22 Jul 1987, R. Hol- zenthal, J. Morse, P. Clausen 12 6, 3 @ (UMSP, INBIO). Rio Peje, 1 km SE San Vicente, 1450 m, 14 Feb 1992, R. Holzen- thal, E Munoz, K. Kjer, 4 2 (INBIO). Re- serva Forestal San Ramon, Rio San Lor- encito, 980 m, 30 Mar 1987, R. Holzenthal, S. Hamilton, M. Heyn, 1 ¢ (UMSP). Guan- acaste: Parque Nacional Guanacaste, Mar- itza, Rio Tempisquito, 550 m, 19 Jul 1987, R. Holzenthal, J. Morse, P Clausen, 12 ¢ (BPS, INBIO, UMSP). Estacion Maritza, Rio Tempisquito, 11 Feb 1990, 2 ¢ (SWRC). Same location, 17 Mar 1989, 1 3,1 2 (BPS). Same location, 11 Apr 1990, 10 3g (SWRC). Same location, 28 May 1989, 4 6, 1 2 (INBIO). Same location, 22 Oct 1990, 6 6, 1 2 (SWRC). Same loca- tion, 11 Nov 1989, 1 3, 1 2 (BPS). Parque Nacional Guanacaste, Rio San Josecito, Es- tacion Mengo, 960 m, 28 Jul 1987, R. Hol- zenthal, J. Morse, P. Clausen, 2 d (UMSP). Parque Nacional Guanacaste, Rio Orosi, Estacion Pitilla, 700 m, 22 May 1990, R. Holzenthal, R. Blahnik, 14 6,5 2 (BPS, UMSP). Parque Nacional Guanacaste, El Hacha, Quebrada Alcornoque, 250 m, 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Jul 1987, R. Holzenthal, J. Morse, P Clau- sen, 1 6 (UMSP). Parque Nacional Rincon de la Vieja, Quebrada Zapilote, 785 m, 3 Mar 1986, R. Holzenthal, 1 6 (BPS). Que- brada Garcia, 470 m, 8 Mar 1986, R. Hol- zenthal, W. Fasth, 1 ¢ (BPS). Comments.—The male syntype is miss- ing but L. Benedetto provided notes on the female syntype. Anacroneuria exquisita, new species Figs. 41—45 Adult habitus.—Head yellow. Median pronotal stripe broad and pale; irregular lat- eral stripes brown, margins pale (Fig. 41). Wings pale amber, transparent circular win- dow beyond cord; veins C, Sc and R pale, others brown. Male.—Forewing length 14 mm. Ham- mer a low membranous disc (Fig. 42). Ae- deagal apex trilobed, lateral lobes small and covered ventrally by membranous process- es. Dorsal keel inconspicuous, hooks slen- der (Figs. 43-45). Female.—Unknown. Nymph.—Unknown. Etymology.—tThe species name, meaning excellent or admirable, refers to the distinc- tive adult color pattern. Types.—Holotype 6 (USNM) and four 3d paratypes (BPS, UMSP) from Costa Rica, San Jose, Parque Nacional Braulio Carrillo, Rio Zurqui, 1650 m, 6 Jul 1986, R. Holzenthal, M. Heyn, B. Armitage. Diagnosis.—A. exquisita is similar in ae- deagal features to A. perplexa (Figs. 88—90) but that species is smaller and darker (Fig. 86). A. ventana has a similar wing pattern but differs in aedeagal structure (Figs. 124— 125), hammer shape (Fig. 122) and head pattern (Fig. 121). A series of female specimens from Es- tacion Maritza are similar in color pattern but they are smaller, have a dark ocellar patch and the dark pronotal stripes are larg- er and convergent posteriorly. Thus these specimens remain unassociated and the fe- male of A. exquisita, unknown. VOLUME 111, NUMBER 3 567 Figs. 41-45. A. exquisita structures. 41. Head and pronotum. 42. Male sternum 9. 43. Aedeagus, lateral. 44. Aedeagus, dorsal. 45. Aedeagus, ventral. Scales: 0.6 mm (41), 0.3 mm (42), 0.15 mm (43-45). 568 Anacroneuria hacha, new species Figs. 48-50, 169-170 Adult habitus.—Dark head pattern covers ocelli and extends to anterior margin; pat- tern diffuse and almost interrupted at M- line. Median pronotal stripe diffuse brown, bordered by yellow; irregular midlateral stripes brown, margins pale (Fig. 46). Wing membrane brown, veins brown. Male.—Forewing length 10-11 mm. Hammer a low indistinct mound (Fig. 47). Aedeagal apex simple, dorsal margin trun- cate or emarginate, keel well developed. Lateral aspect somewhat hatchet shaped; hooks slender (Fig. 48-50, 169-170). Female.—Unknown. Nymph.—Unknown. Etymology.—Hacha, Spanish for axe, re- fering to the hatchet shaped aedeagal apex, is used as a noun in apposition. Types.—Holotype ¢6 (USNM) and 1 ¢ paratype (SWRC) from Costa Rica, Guan- acaste, Estacion Maritza, Rio Tempisquito, 27 Apr 1989. Additional Paratypes: Costa Roca: Type locality, 11 May 1990, 2 6 (SWRC). Same location, 21 May 1990, 1 6 (SWRC). Same location, 30 May 1990, 6 36 (BPS). Same location, 10 Oct 1989, 1 6 (SWRC). Same location, 22 Oct 1990, 4 6 (INBIO). Parque Nacional Guanacaste Maritza, Rio Tempisquito, 550 m, 19 Jul 1987, R. Holzenthal, J. Morse, P. Clausen, 6 36 (UMSP). Same location, 17 Jun 1988, C. M. Flint, O. S. Flint, R. Holzenthal, 10 3 (USNM). Diagnosis.—A. hacha is similar to A. divisa in color pattern (Fig. 36) and general aedeagal shape (Figs. 38—40), but the two are distinguished on the basis of the ham- mer (Figs. 37, 47) and aedeagal keel (Figs. 40, 49). Anacroneuria harperi, new species Figs. 51-55, 142 Anacroneuria sp. B: Harper, 1992:118. Adult habitus.—Head yellow with dif- fuse brown area forward of ocelli; lappets PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON brown. Median pronotal stripe yellow; most of disc pale brown, anterolateral margins pale (Fig. 51). Wing membrane brown, veins brown. Male.—Forewing length 7 mm. Hammer thimble shaped, height greater than diame- ter (Fig. 52). Aedeagal body asymmetrical, apex twisted in a slight sinistral direction. Hooks twisted and asymmetrical; apex of right hook recurved. Dorsal keel absent (Figs. 53-55). Female.—Forewing length 9 mm. Sub- genital plate four lobed; inner lobes smaller than outer lobes. Transverse sclerite of ster- num nine narrow; mesal area of sclerite densely setose. Mesal sclerite T-shaped, lat- eral setae prominent (Fig. 142). Nymph.—Unknown. Etymology.—The patronym honors P. Harper for his recognition of this interesting species. Types.—Holotype 3 from Panama, Ala- juela, 17 Apr 1911, A. Busck (USNM). Paratypes: Panama: Alajuela: 6 Apr 1911, A. Busck, 1 36, 1 2 (USNM). Same loca- tion, 16 Apr 1911, A. Busck, 1 6 (USNM). Same location, 8 Mar 1912, A. Busck, 1 2 (USNM). Darien Province: Santa Fe, 28 May 1967, D. M. Delong, C. A. Triplehorn, 1 ¢ (OSU). Diagnosis.—In size and general colora- tion this species is similar to A. curiosa and A. uatsi but the twisted aedeagal apex (Figs. 53-55) is unique to this species. Anacroneuria holzenthali, new species Figs. 56-60, 143, 171-172 Adult habitus.—Dark pattern covers ocelli and extends forward to midpoint of lappets; pattern interrupted by a pair of minute pore sized unpigmented spots and a larger mesal spot. Pronotum dark except for slender mesal stripe and anterolateral mar- ginal spots (Fig. 56). Wing membrane brown, veins brown. Male.—Forewing length 10-12 mm. Hammer thimble shaped (Fig. 57). Aedea- gal apex multilobed, shoulders winged. VOLUME 111, NUMBER 3 569 Figs. 46-50. A. hacha structures. 46. Head and pronotum. 47. Male sternum 9. 48. Aedeagus, lateral. 49. Aedeagus, dorsal. 50. Aedeagus, ventral. Scales: 0.6 mm (46), 0.3 mm (47), 0.15 mm (48-50). 570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 51-55. A. harperi structures. 51. Head and pronotum. 52. Male sternum 9. 53. Aedeagus, ventral. 54. Aedeagus, lateral. 55. Aedeagus, dorsal. Scales: 0.6 mm (51), 0.3 mm (52), 0.15 mm (53-55). VOLUME 111, NUMBER 3 571 Figs. 56-60. A. holzenthali structures. 56. Head and pronotum. 57. Male sternum 9. 58. Aedeagus, lateral. 59. Aedeagus, dorsal. 60. Aedeagus, ventral. Scales: 0.6 mm (56), 0.3 mm (57), 0.15 mm (58-60). 572 Dorsal keel transverse, hooks slender (Figs. 58-60, 171-172). Female.—Forewing length 14-15 mm. Subgenital plate four lobed; mesal notch deep, V-shaped, lateral notches shallow. Transverse sclerite of sternum nine sinuate, mesal sclerite T-shaped and setose. Setae on sclerite stalk minute, lateral setae prominent (Fig. 143). Nymph.—Unknown. Etymology.—The patronym honors R. W. Holzenthal for his outstanding contributions to our understanding of the Costa Rican aquatic insect fauna. Types.—Holotype 3 from Costa Rica, Guanacaste, Parque Nacional Rincon de la Vieja, Quebrada Zapilote, 785 m, 3 Mar 1986, R. Holzenthal (USNM). Paratypes: Costa Rica: Alajuela, Reserva Forestal San Ramon, Rio San Lorencito, 980 m, 6—10 Mar 1991, R. Holzenthal, E Munoz, J. Huisman, 6 36, 11 2 (INBIO, UMSP). Guanacaste: Quebrada Garcia, 10.6 km ENE Quebrada Grande, 470 m, 8 Mar 1986, R. Holzenthal, W. Fasth, 3 6 (UMSP). Guanacaste, Parque Nacional Guanacaste, Maritza, Rio Tempisquito, 550 m, 17 Jun 1988, C. M. Flint, O. S. Flint, R. Holzenthal, 4 6, 1 2 (USNM). Same lo- cation, 19 Jul 1987, R. Holzenthal, J. Morse, P. Clausen, 2 6 (UMSP). Parque Nacional Guanacaste, Rio Tempisquito Sur, Maritza, 600 m, 30 Aug 1990, J. Huisman, E Quesada, 1 ¢, 1 2 (BPS). Estacion Mar- itza, Rio Tempisquito, 11 May 1990, 2 d (SWRC). Same location, 17 Mar 1989, 1 3,1 2 mating (SWRC). Same location, 28 Apr 1989, 1 6, 1 2 (SWRC). Heredia: Quebrada Chiquiza, Rava Avis Biological Station, 550 m, 31 Mar 1989, R. Blahnik, Solis 1 ¢ (UMSP). Nicaragua: Chontales, Santo Domingo, Nov 1985, J. M. Maes, 1 36 (USNM). Diagnosis.—See under A. benedettoi. Anacroneuria lineata (Navas) Figs. 61-65 Neoperla lineata Navas, 1924:73. Holotype 3, Costa Rica (MNHN). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Adult habitus.—Head with dark pattern over ocelli extending to M-line; lappets and small triangular patch forward of M-line dark. Median pronotal stripe diffuse brown, bordered by yellow; margins dark (Fig. 61). Wing membrane transparent, veins brown. Male.—Forewing length 12 mm. Ham- mer nipple shaped (Fig. 62). Aedeagal apex broad, scoop shaped with a pair of small lateral projections at shoulder. Dorsal keel prominent, hooks slender (Figs. 63-65). Female.—Unknown. Nymph.—Unknown. Material.—Costa Rica: Alajuela: Rio Pi- zote, ca. 5 km N Dos Rios, 470 m, 9 Mar 1986, R. Holzenthal, W. Fasth, 1 6 (UMSP). Parque nacional Rincon de la Vie- ja, Quebrada Provision, 810 m, 4 Mar 1986, R. Holzenthal, W. Fasth, 1 6 (UMSP). Guanacaste: Rio Mena, 4.2 km W Santa Cecilia, 11 Mar 1986, R. Holzenthal, W. Fasth, 1 6 (UMSP). Parque Nacional Guanacaste, Maritza, Rio Tempisquito, 550 m, 17 Jun 1988, C. M. Flint, O. S. Flint, R. Holzenthal, 3 6 (USNM). Estacion Marit- za, Rio Tempisquito, 10 Oct 1989, 1 6 (SWRC). Same location, 11 Apr 1990, 1 6 (SWRC). Same location, 11 May 1990, 2 3d (INBIO). Rio Gongora, 4 km NE Que- brada Grande, 590 m, 21 Jul 1987, R. Hol- zenthal, J. Morse, P Clausen, 10 6 (UMSP, BPS). San Jose: Palmital de Guarco, Cerro de la Muerte, km 37 Pan American High- way, 7 May 1995, R. W. Baumann, B. Houseman, 1 6 (BYU). MEXICO: Chia- pas: 10 mi N Palenque, 14 June 1971, J. Zimmerman, 6 6,5 2 (BPS). Comments.—Figures of the holotype ae- deagus provided by L. Benedetto seem to match this species. Anacroneuria magnirufa Jewett Figs. 66—70, 144, 155, 173-174 Anacroneuria magnirufa Jewett, 1958:162. Holotype 2, El Volcan, Chiriqui, Panama (AMNH). Adult habitus.—Dark brown pigment ex- tends from behind ocelli to M-line; lateral VOLUME 111, NUMBER 3 573 63 65 Figs. 61-65. A. Jineata structures. 61. Head and pronotum. 62. Male sternum 9. 63. Aedeagus, lateral. 64. Aedeagus, dorsal. 65. Aedeagus, ventral. Scales: 0.6 mm (61), 0.3 mm (62), 0.15 mm (63-65). 64 574 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON UU A Gy Ie LORE “Nt Uc ends Memne COHAN ens iia lar ce) Figs. 66-70. A. magnirufa structures. 66. Head and pronotum. 67. Male sternum 9. 68. Aedeagus, lateral. 69. Aedeagus, dorsal. 70. Aedeagus, ventral. Scales: 0.6 mm (66), 0.3 mm (67), 0.15 mm (68-70). VOLUME 111, NUMBER 3 callosities prominent; lappets and mesal field forward of M-line dark brown. Pro- notum brown except for scattered pale ru- gosities and a narrow pale mesal band (Fig. 66). Wing membrane and veins dark brown. Male.—Forewing length 17—20 mm. Hammer absent (Fig. 67). Aedeagal apex narrow and scoop shaped, ventral aspect with a pair of small membranous processes. Dorsal keel a thin transverse ridge, hooks slender (Figs. 68—70, 173-174). Female.—Forewing length 23-26 mm. Subgenital plate bilobed, margins truncate or slightly rounded. Mesal notch shallow, small longitudinal tubercle forward of notch. Transverse sclerite absent from ster- num nine, mesal sclerite sparsely setose (Fig. 144). Nymph.—Unknown. Material.—Costa Rica: Alajuela: Rio Peje, 1 km SE San Vicente, 1450 m, 15 Feb 1992, R. Holzenthal, EK Munoz, K. Kjer, 26 3, 15 2 (BPS, INBIO, UMSP). Rio Pizote, 5 km S Brasilia, 390 m, 12 Mar 1986, R. Holzenthal, W. Fasth, 2 6 (UMSP). Rio Bochinche, Cerro Campana, 600 m, 22 Jul 1987, R. Holzenthal, J. Morse, P. Clausen, 1 ¢ (UMSP). Rio San Lorencito, Reserva Forestal San Ramon, 980 m, 1 Apr 1987, R. Holzenthal, S. Hamilton, M. Heyn, 1 ¢, 2 2. Same location, 6-10 Mar 1991,.R. Holzenthal, EF Munoz, J. Huisman, 3 6 (BPS). Guanacaste: Rio Los Ahogados, 11.3 km ENE Quebrada Grande, 470 m, 7 Mar 1986, R. Holzenthal, W. Fasth, 2 d, 1 2 (UMSP). Rio Gongora, 590 m, 21 Aug 1987, R. Holzenthal, J. Morse, P. Clausen, 1 36, 1 2 (UMSP). Estacion Maritza, Rio Tempisquito, 14 Jan 1990, 1 56 (SWRC). Same location, 11 Feb 1990, 1 ¢ (SWRC). Same location, 11 Apr 1990, 1 ¢ (SWRC). Same location, 28 Apr 1989, 1 ¢6, 3 (SWRC). Same location, 11 May 1990, 1 3 (SWRC). Same location, 30 May 1990, 1 5 (SWRC). Same location, 5 Jul 1989, 1 2 (SWRC). Same location, 19 Aug 1989, 2 6,2 2 (SWRC). Same location, 10 Oct 1989, 2 6, 1 2 (SWRC). Same location, 11 Nov 1989, 1 ¢ (SWRC). Puntarenas: Rio 575 Bellavista, ca. 1.5 km NW Las Alturas, 1400 m, 10 Aug 1990, R. Holzenthal, R. Blahnik, EK Munoz, 2 6 (UMSP). Same lo- cation, 16 Mar 1991, R. Holzenthal, KF Mu- noz, J. Huisman, 3 ¢ (BPS). Panama: Chi- riqui, Guadalupe Arriba, 11-17 Sep 1985, H. Wolda, 2 6 (BYU). Same location, 10— 16 Jul 1985, H. Wolda, 2 ¢d (BYU). Same location, 24—30 Jul 1985, H. Wolda, 1 ¢ (BYU). Comments.—Collections of A. magnirufa in Costa Rica are known from throughout the year, and the known species range is from Panama to Honduras. Few specimens are available from Panama and these are larger and have a slightly wider aedeagal apex and less conspicuous dorsal keel than Costa Rican specimens. No specimens are known from Nicaragua and only two para- type males are known from Honduras. All Costa Rican female specimens have a mesal tubercle near the subgenital plate notch. This conspicuous feature was not men- tioned by Jewett (1958) and is not shown in figures of the holotype made by Bene- detto (L. Benedetto, pers. comm.). This variation in both male and female speci- mens from Chiriqui suggests the possibility that two closely related species are in- volved. Anacroneuria marca, new species Figs. 71-75, 145, 175-176 Adult habitus.—Head with a pair of brown L-shaped marks extending from ocelli but sometimes fused between ocelli; lappets brown. Pronotum with a narrow median yellow band; lateral area brown but with scattered pale rugosities and pale an- terolateral margin (Fig. 71). Wing mem- brane transparent, veins brown. Male.—Forewing length 16 mm. Ham- mer thimble shaped, height subequal to ba- sal diameter (Fig. 72). Aedeagal apex sim- ple, abruptly narrowed from shoulders to a slender scoop. Hooks slender, dorsal keel absent (Figs. 73—75, 175-176). Female.—Forewing length 20 mm. 576 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 71-75. A. marca structures. 71. Head and pronotum. 72. Male sternum 9. 73. Aedeagus, lateral. 74. Aedeagus, dorsal. 75. Aedeagus, ventral. Scales: 0.6 mm (71), 0.3 mm (72), 0.15 mm (73-75). VOLUME 111, NUMBER 3 577 Sa 79 BO Figs. 76-80. A. marginata structures. 76. Head and pronotum. 77. Male sternum 9. 78. Aedeagus, lateral. 79. Aedeagus, dorsal. 80. Aedeagus, ventral. Scales: 0.6 mm (76), 0.3 mm (77), 0.15 mm (78-80). Subgenital plate bilobed; notch V-shaped, chia; setal patch of mesal sclerite inter- lobes truncate to emarginate. Transverse rupted into a median stalk and lateral setal sclerite of sternum nine wide; interseg- areas (Fig. 145). mental membrane covered with microtri- Nymph.—Unknown. 578 Etymology.—Marca, Spanish for brand, refers to the L-shaped marks on the head, and is used as a noun in apposition. Types.—Holotype ¢ from Costa Rica, Heredia, Parque Nacional Braulio Carrillo, Rio Peje, 480 m, 29 May 1990, R. Holzen- thal, R. Blahnik, E Munoz (USNM). Para- types: Costa Rica: Alajuela: Rio Peje, ca. 1 km SE San Vicente, 1450 m, 14 Feb 1992, R. Holzenthal, EF Munoz, K. Kjer, 3 6 (UMSP). Reserva Forestal San Ramon, Rio San Lorencito, 980 m, 30 Mar 1987, R. Holzenthal, S. Hamilton, M. Heyn, 9 @ (UMSP). Same location, 6 Mar 1991, R. Holzenthal, EF Munoz, J. Huisman, 3 2 (IN- BIO). Cartago: Reserva Tapanti, Rio Gran- de de Orosi, 1650 m, 8 Jul 1986, R. Hol- zenthal, M. Heyn, B. Armitage, 3 6, 3 @ (BPS). Puntarenas: Rio Coton, Las Alturas, 1360 m, 18 Mar 1991, R. Holzenthal, FE Munoz, J. Huisman, 1 6, 1 2 (INBIO). Same location, 16 Feb 1986, R. Holzenthal, J. Morse, W. Fasth, 1 ¢ (BPS). Zona Pro- tectora Las Tablas, Rio Coton, Sitio Coton, 1460 m, 15 Apr 1989, R. Holzenthal, R. Blahnik, 1 d6, 1 2 (USNM). San Jose: Rio Chirripo Pacifica, 9.5 km NE Rivas, 1370 m, 23 Feb 1986, R. Holzenthal, J. Morse, W. Fasth, 1 2 (UMSP). Diagnosis.—The color pattern of A. mar- ca is similar to that of A. ventana (Fig. 121) but hammer shape (Fig. 122) and the ae- deagal keel (Fig. 124) distinguish these spe- cies. A. zapata is also similar but has a tri- lobed aedeagal apex (Fig. 130). Anacroneuria marginata, new species Figs. 76-80, 177-178 Adult habitus.—Diffuse brown pattern extends from ocelli to M-line and laterally to lappets. Pronotum with narrow pale me- dian and marginal stripes; dark midlateral stripes rugose (Fig. 76). Wing membrane amber, veins brown. Male.—Forewing length 8 mm. Hammer thimble shaped, height less than basal di- ameter (Fig. 77). Aedeagal apex simple, broad and rounded at tip. Dorsal keel mod- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON erately developed, hooks short (Figs. 78— 80, 177-178). Female.—Unknown. Nymph.—Unknown. Etymology.—The species name, meaning bordered, refers to the pale marginal bands on the pronotum. Types.—Holotype 6 from Costa Rica, Guanacaste, Parque Nacional Guanacaste, Maritza, Rio Tempisquito, 550 m, 17 Jun 1988, C. M. Flint, O. S. Flint, R. Holzenthal (USNM). Paratypes: Costa Rica: Guana- caste: Parque Nacional Guanacaste, Marit- za, Rio Tempisquito Sur, 30 Aug 1990, 600 m, J. Huisman, E Quesada, 1 ¢ (UMSP). Estacion Maritza, Rio Tempisquito, 14 Jan 1990, 1 6 (SWRC). Same location, 17 Mar 1989, 1 d (SWRC). Same location, 27 Apr 1989, 2 3 (BPS). Same location, 28 May 1989, 1 6 (SWRC). Same location 30 May 1990, 1 6 (INBIO). Same location, 22 Oct 1990, 1 3d (INBIO). Diagnosis.—The aedeagus of this species is similar to that of A. costana (Figs. 28— 30) but the apex is shorter and wider and the keel more pronounced in A. marginata. Anacroneuria maritza, new species Figs. 81-85, 146, 151—152 Adult habitus.—Dark brown covers ocel- li and extends forward to M-line; lappets and a small mesal spot forward of M-line brown. Median brown pronotal stripe bor- dered by yellow; irregular midlateral stripes brown, margins pale (Fig. 81). Wing mem- brane brown, veins brown except pale C and Sc. Male.—Forewing length 12-13 mm. Hammer thimble shaped, height less than basal diameter (Fig. 82). Aedeagal apex scoop shaped, tip with parallel margins and well developed dorsal keel. Ventral aspect covered with large membranous lobes; hooks slender (Figs. 83-85). Female.—Forewing length 15-16 mm. Subgenital plate bilobed, notch broadly V- shaped. Transverse sclerite of sternum nine absent; mesal sclerite V-shaped and sparse- VOLUME 111, NUMBER 3 579 Capone 4° Figs. 81-85. Zi A. maritza structures. 81. Head and pronotum. 82. Male sternum 9. 83. Aedeagus, lateral. 84. Aedeagus, dorsal. 85. Aedeagus, ventral. Scales: 0.6 mm (81), 0.3 mm (82), 0.15 mm (83-85). 580 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 86-90. A. perplexa structures. 86. Head and pronotum. 87. Male sternum 9. 88. Aedeagus, lateral. 89. Aedeagus, dorsal. 90. Aedeagus, ventral. Scales: 0.6 mm (86), 0.3 mm (87), 0.15 mm (88—90). ly setose. Patch of long red-brown setae at oval areas anterolateral to ocelli. Pronotum base of lateral bar (Fig. 146). brown with scattered pale rugosities (Fig. Nymph.—Body length 14-17 mm. Head _ 151). pattern brown except for pale M-line and Etymology.—The species name, based on VOLUME 111, NUMBER 3 581 ig: Pie, ae Figs. 91-95. A. planicollis structures. 91. Head and pronotum. 92. Male sternum 9. 93. Aedeagus, lateral. 94. Aedeagus, dorsal. 95. Aedeagus, ventral. Scales: 0.6 mm (91), 0.3 mm (92), 0.15 mm (93-95). the type locality, is used as a noun in ap- quito, 11 Feb 1989 (USNM). Paratypes: position. Costa Rica: Same location, 16 Mar 1989, 1 Types.—Holotype 3 from Costa Rica, od (BPS). Same location 17 Mar 1989, 1 3 Guanacaste, Estacion Maritza, Rio Tempis- (INBIO). Same location, 7 Apr 1989, 3 d, 582 2 2 (SWRC, BPS). Same location, 11 Apr 1990, 1 ¢6 (INBIO). Same location, 27 Apr 1989, 1 d (SWRC). Same location, 24 May 1989, 3 3 (BPS, SWRC). Same location, 28 May 1989, 1 2 (USNM). Same location, 22 Oct 1990, 1 6 (SWRC). Same location, 11 Nov 1989, 1 ¢ (INBIO). Same location, reared from eggs, 2 6, 2 2 (SWRC). Same location, 30 Aug 1990, J. Huisman, R. Blahnik, EF Quesada, 1 6 (UMSP). Guan- acaste: Rio Los Ahogados, 470 m, 7 Mar 1986, R. Holzenthal, W. Fasth, 1 o (UMSP). Hacienda la Pacifica, 75 m, 8 Jun 1973, G. Ekis, 1 d (BYU). Monteverde, 22 Jun 1986, W. Hanson, 7 6 (USU). Diagnosis.—The color pattern is similar to several species including A. divisa (Fig. 36) and A. planicollis (Fig. 91) but aedeagal features (Figs. 39, 85, 95) easily distinguish these species. Comments.—Scientists at the Stroud Wa- ter Research Center reared two males and two females of this species from eggs col- lected at Estacion Maritza (J. Jackson, pers. comm.). The first egg mass collected 26 Aug 1992 produced a female on 18 Feb 1993 and a male on 11 Mar 1993. The sec- ond egg mass from 29 Nov 1992 produced a male on 14 May 1994 and a female on 3 Jun 1994. The nymphal description is based on exuviae from these specimens. Anacroneuria perplexa, new species Figs. 86-90, 179-180 Adult habitus.—Dark head pattern covers ocelli and extends anterolaterally to lappets; small pale spot located in center of head. Pronotum mostly brown but with incon- spicuous mesal yellow band (Fig. 86). Wing membrane brown, veins brown. Male.—Forewing length 8 mm. Hammer absent or reduced to obscure membranous spot (Fig. 87). Aedeagal apex trilobed, lat- eral lobes small and ear shaped. Dorsal keel well developed, hooks enlarged and some- what chelate (Figs. 88-90, 179-180). Female.—Unknown. Nymph.—Unknown. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Etymology.—The species name, meaning tangled or confused, refers to the initial dif- ficulty experienced in separating this spe- cies from other small, dark forms with poorly developed hammers. Types.—Holotype ¢ (USNM) and 2 6 paratypes (UMSP) from Costa Rica, Limon, Rio Barbilla, ca. 8 km W B-line, 30 m, 31 Jan 1986, R. Holzenthal, J. Morse, W. Fasth. Additional paratypes: Costa Rica: Li- mon: Reserva Biol. Hitoy-Cerere, Rio Cer- ere, 90 m, 23 Mar 1987, R. Holzenthal, S. Hamilton, M. Heyn, 8 d (UMSP, BPS). Rio Banano, 16 km WSW Bomba, 150 m, 26 Mar 1987, R. Holzenthal, S. Hamilton, M. Heyn, (UMSP). Puntarenas: Rio Jaba, 1.4 km W Las Cruces, 1150 m, 14 Jun 1986, R. Holzenthal, M. Heyn, B. Armitage, 1 ¢ (INBIO). Villa Neilly, 21 Jun 1964, R. Woodruff, 1 6 (FSCA). San Jose: Escazu, 7-10 Apr 1988, FE D. Parker, 1 ¢ (USU). Diagnosis.—A male paratype of A. cren- ulata from Temescaltepec, Mexico, is very similar to this species and distinct from the allotype of A. crenulata. The Mexican spec- imen differs from Costa Rican specimens most conspicuously in having long slender aedeagal hooks. Additional comments are given under A. exquisita. Anacroneuria planicollis Klapalek Figs. 91-95, 147 Anacroneuria planicollis Klapalek, 1923: 22. Lectotype 3, Sierra du Naya, Mexico (NMP). Anacroneuria dampfi Jewett, 1958:165. New synonymy. Anacroneuria chiapasa Jewett, 1958:166. New synonymy. Adult habitus.—Head pattern usually with brown ocellar spot, shading to diffuse brown forward of ocelli; lappets brown. Median pronotal stripe brown, bordered by yellow; lateral brown stripes interrupted by scattered pale areas on disc and along mar- gins (Fig. 91). Wing membrane transparent, veins brown except C and Sc pale. Male.—Forewing length 14-15 mm. VOLUME 111, NUMBER 3 Hammer thimble shaped, height subequal to basal diameter (Fig. 92). Aedeagal apex tri- lobed, mesal lobe slightly larger. Hooks slender, dorsal keel well developed (Figs. 93-95). Female.—Forewing length 16-17 mm. Subgenital plate bilobed, lobes truncate to emarginate. Mesal sclerite of sternum nine T-shaped; lateral arms with long setae, stalk with fine short setae (Fig. 147). Nymph.—Unknown. Material.—Costa Rica; Monteverde, 22 Jun 1986, W. J. Hanson, 2 ¢ (USU). Ala- juela: 20 km S Upala, 12 Feb—5 Mar 1991, FE Parker, 2 6 (USU). Puntarenas: Rio Bel- lavista, ca. 1.5 km NW Las Alturas, 1400 m, 15 Jun 1986, R. Holzenthal, M. Heyn, B. Armitage, 1 ¢ (UMSP). Same location, 8 Apr 1987, R. Holzenthal, S. Hamilton, M. Heyn, 1 6, 2 2 (INBIO). Mexico: Sierra du Naya, L. Diguet, 1898, lectotype 6, par- alectotype 2 (NMP). Nicaragua: Nuevose- govia, Dipilto, Dec 1986, J. Maes, 1 6 (USNM). Panama: Chiriqui, Cerro Punta, 3 mi N on Pan American Highway, 4 Apr 1954, G. Field, 1 d (USNM). Comments.—The type series included four male and three female specimens from Mexico and a female from Costa Rica (Kla- palek 1923). Of these, a syntype male and female were found in the Natural History Museum, Prague and studied while on loan to P. Zwick. the male is designated as lec- totype. Synonymy of the two Jewett (1958) species is based on study of the holotypes and male allotypes. Anacroneuria plutonis (Banks) Figs. 96-100, 148, 157, 181-182 Neoperla plutonis Banks, 1914:610. Lec- totype 2, La Trinidad, Costa Rica (MCZ). Forquilla_ tristani Navas, synonymy. 1932:29. New Adult habitus.—Head pattern dark brown from ocellar area to M-line except for pale oval spots lateral to ocelli; lappets connect- ed by a diffuse brown band along anterior 583 margins of M-line. Pronotum diffuse brown to dark brown but without distinct bands (Fig. 96). Wing membrane brown, veins brown. Male.—Forewing length 19-21 mm. Hammer thimble shaped, height less than basal diameter (Fig. 97). Aedeagal apex simple, gradually tapered to a narrow tip. Hooks slender, dorsal keel absent (Figs. 98— 100, 157, 181-182). Female.—Forewing length 25-28 mm. Subgenital plate bilobed; lobes truncate, notch shallow and U-shaped. Transverse sclerite absent, mesal sclerite T-shaped, sparsely setose (Fig. 148). Nymph.—Unknown. Material.—Costa Rica: Alajuela: Rio Sarapiqui, ca. 2 km SE Cariblanco, 710 m, 22 Jun 1986, R. Holzenthal, M. Heyn, B. Armitage, 1 ¢d, 1 2 (BPS). Quebrada Latas, 8.9 km NE Bajos del Toro, 1030 m, 6 Sep 1990, R. Holzenthal, R. Blahnik, J. Huis- man, 2 6 (UMSP). Reserva Forestal San Ramon, Rio San Lorencito, 6—10 Mar 1991, R. Holzenthal, EF Munoz, J. Huisman, 4 d (UMSP, INBIO). Cartago: Reserva Tapanti, Quebrada Palmitos, 1400 m, 24 Mar 1991, R. Holzenthal, EF Munoz, J. Huisman, 1 2 (UMSP). Same location, 2 Jun 1990, R. Holzenthal, R. Blahnik, FEF Munoz, 7 ¢ (BPS, INBIO). Guanacaste: Estacion Mar- itza, Rio Tempisquito, 14 May 1989, 1 ¢ (SWRC). Parque Nacional Guanacaste, Maritza, Rio Tempisquito, 550 m, 19 Jul 1987, R. Holzenthal, J. Morse, P. Clausen, 1 36 (UMSP). Puntarenas: Rio Bellavista, ca. 1.5 km NW Las Alturas, 1400 m, 8 Apr 1987, R. Holzenthal, S. Hamilton, M. Heyn, 1 6 (UMSP). Rio Coton, Las Alturas, 1360 m, 16 Feb 1986, R. Holzenthal, J. Morse, W. Fasth, 1 6 (UMSP). Same location, 18 Mar 1991, R. Holzenthal, E Munoz, J. Huisman, | ¢ (BPS). San Jose: El Salvaje, Rio Tabarcia, 8 km E Palmichal, 1650 m, 19 Jan 1992, R. Holzenthal, K. Kjer, E Quesada, 11 6,3 2 (BPS, UMSP, INBIO). Rio Parrita Chiquita, 1990 m, 18 Jun 1986, R. Holzenthal, M. Heyn, B. Armitage, 1 6, 1 2 (USNM). 584 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON - 98 Figs. 96-100. A. plutonis structures. 96. Head and pronotum. 97. Male sternum 9. 98. Aedeagus, lateral. 99. Aedeagus, dorsal. 100. Aedeagus, ventral. Scales: 0.6 mm (96), 0.3 mm (97), 0.15 mm (98-100). Comments.—Anacroneuria plutonis and A. tristani are both large dark species described from Costa Rican specimens. Navas (1932) reports the forewing length of A. tristani fe- males is 18.5 mm but the type specimen is actually larger than the A. plutonis lectotype from La Trinidad (L. Benedetto, pers. comm.). I have not seen the specimens iden- tified by Jewett (1958) as A. tristani but the four lobed female subgenital plate mentioned VOLUME 111, NUMBER 3 in Jewett’s description clearly indicates this material was misidentified. Anacroneuria talamanca, new species Figs. 101-105, 149 Adult habitus.—Ocellar area covered by a small brown quadrangle; lateral margins of frons diffuse brown, lappets brown. Me- dian pronotal stripe yellow, lateral stripes brown (Fig. 101). Wing membrane trans- parent veins brown. Male.—Forewing length 7—8 mm. Ham- mer thimble shaped, height subequal to ba- sal diameter (Fig. 102). Aedeagal apex sim- ple, slender with small dorsal keel. Hooks enlarged and ventrally keeled (Figs. 103— 105). Female.—Forewing length 9-10 mm. Subgenital plate bilobed; lobes truncate, median notch shallow and V-shaped. Trans- verse sclerite of sternum nine poorly de- veloped, mesal sclerite with long stem, lat- eral arms poorly developed and sparsely se- tose (Fig. 149). Nymph.—Unknown. Etymology.—The species name, based on the Cordillera Talamanca, is used as a noun in apposition. Types.—Holotype ¢ from Costa Rica, San Jose, Reserva Biol. Carara, Rio Carara, Carara, 200 m, 14 Mar 1991, R. Holzenthal, E Munoz, J. Huisman (USNM). Paratypes: Costa Rica: Heredia: Estacion Biol. La Sel- va, Rio Puerto Viejo, 30 m, 10 Feb 1986, R. Holzenthal, 1 6 (UMSP). La Selva, 14 Jun 1986, W. J. Hanson, 1 d (USU). Pan- ama: Bocas del Toro: Miramar, 1 May 1979, H. Wolda, 1 6, 2 2 (BYU). Same location, 22 May 1979, H. Wolda, 1 & (BYU). Diagnosis.—The aedeagus of this species is similar to that of the allotype of A. cren- ulata Jewett and to the holotype of A. litura (Pictet) (Zwick 1972). A. talamanca differs from these in having the apical aedeagal section beyond the hooks about as long as wide; in A. crenulata and A. litura this sec- tion is distinctly longer than wide and has 585 a dorsal keel more than twice the length of the keel of A. talamanca. In addition, the female holotype of A. crenulata has a four lobed subgenital plate whereas the subgen- ital plate of A. talamanca is bilobed. Anacroneuria tornada, new species Figs. 106-110 Adult habitus.—Diffuse brown area for- ward of ocelli, lappets brown. Narrow brown median pronotal stripe bordered by narrow yellow stripes; midlateral stripes brown, margins pale (Fig. 106). Wing membrane transparent, veins brown. Male.—Forewing length 16 mm. Ham- mer a low disc (Fig. 107). Aedeagal apex multilobed, curved abruptly ventrad be- tween hook tips. Apex in dorsal aspect somewhat truncate, keel absent. Hooks stout, enlarged apically (Figs. 109-110). Female.—Unknown. Nymph.—Unknown. Etymology.—Tomada, Spanish for rene- gade, refers to the unusual aedeagal apex of this species and is used as a noun in ap- position. Types.—Holotype ¢ from Costa Rica, Cartago, Reserva Tapanti, Quebrada Pal- mitos, 1400 m, 2 Jun 1990, R. Holzenthal, R. Blahnik, E Munoz (USNM). Diagnosis.—The color pattern of A. tor- nado is similar to that of A. maritza (Fig. 81), A. divisa (Fig. 36) and A. planicollis (Fig. 51) but the aedeagal apex clearly dis- tinguishes these species (Figs. 39, 85, 95, 110). Anacroneuria uatsi, new species Figs. 111-115, 150, 153-154 Adult habitus.—Head yellow except dark areas on lappets and over ocelli. Median third of pronotum pale, lateral margins brown (Fig. 111). Wing membrane trans- parent, veins brown except pale C and Sc. Male.—Forewing length 9 mm. Hammer thimble shaped, length greater than basal diameter (Fig. 112). Aedeagal apex a slen- der, finger shaped structure arising from a 586 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 103. 104. +~—«:105 Figs. 101-105. A. talamanca structures. 101. Head and pronotum. 102. Male sternum 9. 103. Aedeagus, lateral. 104. Aedeagus, dorsal. 105. Aedeagus, ventral. Scales: 0.6 mm (101), 0.3 mm (102), 0.15 mm (103-— 105). VOLUME 111, NUMBER 3 587 Figs. 106-110. A. tornada structures. 106. Head and pronotum. 107. Male sternum 9. 108. Aedeagus, lateral. 109. Aedeagus, dorsal. 110. Aedeagus, ventral. Scales: 0.6 mm (106), 0.3 mm (107), 0.15 mm (108-110). 588 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 114 115 Figs. 111-115. A. watsi structures. 111. Head and pronotum. 112. Male sternum 9. 113. Aedeagus, lateral. 114. Aedeagus, dorsal. 115. Aedeagus, ventral. Scales: 0.6 mm (111), 0.3 mm (112), 0.15 mm (113-115). broad base with rounded, slightly projecting Transverse sclerite of sternum nine narrow; shoulders. Dorsomesal keel minute, trian- mesal T-shaped sclerite setose (Fig. 150). gular, hooks slender (Figs. 113-115). Nymph.—Body length 9-11 mm. Head Female.—Forewing length 10 mm. Sub- forward of ocelli dark brown except for in- genital plate with four subequal lobes. complete M-line and anterior margin. Pro- VOLUME 111, NUMBER 3 589 Figs. 116-120. A. varilla structures. 116. Head and pronotum. 117. Male sternum 9. 118. Aedeagus, lateral. 119. Aedeagus, dorsal. 120. Aedeagus, ventral. Scales: 0.6 mm (116), 0.3 mm (117), 0.15 mm (118-120). 590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Nee) LES oan: PT a a TS Figs. 121-125. A. ventana structures. 121. Head and pronotum. 122. Male sternum 9. 123. Aedeagus, lateral. 124. Aedeagus, dorsal. 125. Aedeagus, ventral. Scales: 0.6 mm (121), 0.3 mm (122), 0.15 mm (123-125). notum predominantly brown (Fig. 153). the type locality, is used as a noun in ap- Anterodorsal femoral bristles grouped in ir- _ position. regular basal and apical patches and a me- Types.—Holotype 6 and ¢@ paratype dian transverse row (Fig. 154). from Costa Rica, Limon, Rio Uatsi, 8 km Etymology.—The species name, basedon W Bribri, 60 m, 25 Mar 1987, R. Holzen- VOLUME 111, NUMBER 3 591 126 aaa eee Figs. 126-130. A. zapata structures. 126. Head and pronotum. 127. Male sternum 9. 128. Aedeagus, lateral. 129. Aedeagus, dorsal. 130. Aedeagus, ventral. Scales: 0.6 mm (126), 0.3 mm (127), 0.15 mm (128-130). 592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 131-135. A. zarpa structures. 131. Head and pronotum. 132. Male sternum 9. 133. Aedeagus, lateral. 134. Aedeagus, dorsal. 135. Aedeagus, ventral. Scales: 0.6 mm (131), 0.3 mm (132), 0.15 mm (133-135). VOLUME 111, NUMBER 3 sI9)3) Fi pesca ats pes Cl t. 31 oi bid 7M Figs. 136-141. Anacroneuria female sterna 8 and 9. 136. A. acutipennis. 137. A. annulipalpis. 138. A. benedettoi. 139. A. blanda. 140. A. varilla. 141. A. divisa. Scale = 0.6 mm. thal, S. Hamilton, M. Heyn (USNM). Ad- (UMSP). Las Canas, Rio Carabici, 26 Jul ditional paratypes: Costa Rica: Guanacaste: 1967, O. S. Flint, 1 gd (USNM). Limon: Rio Tempisquito, 3 km S route 1,75 m,6 E.A.R.T-H., Rio Destierra, Pozo Azul, 15 Mar 1986, R. Holzenthal, W. Fasth, 2 ¢ m, 5 Feb 1992, R. Holzenthal, EF Munoz, 594 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 142-147. pfs “ETT. 7 oh! Se ite of ‘ DE ipa ye statd 2 Que mete 147 Anacroneuria female sterna 8 and 9. 142. A. harperi. 143. A. holzenthali. 144. A. magnirufa. 145. A. marca. 146. A. maritza. 147. A. planicollis. Scale = 0.6 mm. K. Kjer, 1 6, 1 6 (BPS). Reserva Biol. Hi- toy-Cerere, Rio Cerere, 90 m, 23 Mar 1987, R. Holzenthal, S. Hamilton, M. Heyn, 1 3d (UMSP). Rio Uatsi, W Uatsi, 50 m, 2 Feb 1986, R. Holzenthal, J. Morse, W. Fasth, 2 nymphs (UMSP). Puntarenas: 2.8 mi E Golfito, 18 Jul 1967, O. S. Flint, 1 6 (USNM). Rio Rincon, 6.5 km S Rincon, 20 m, 7 Apr 1987, R. Holzenthal, S. Hamilton, M. Heyn, 1 6 (INBIO). Quebrada Pita, 3 km W Golfito, 15 m, 15 Feb 1986, R. Hol- zenthal, J. Morse, W. Fasth, 22 2 (UMSP, INBIO). Quebrada Portera, N Grande Por- tera, 5 Jul 1992, T. Shepard, 1 ¢d (GSU). Diagnosis.—See under A. curiosa. The nymphs were associated by dissection of male genitalia from a pre-emergent individ- ual. VOLUME 111, NUMBER 3 595 Figs. 148-150. Anacroneuria female sterna 8 and 9. 148. A. plutonis. 149. A. talamanca. 150. A. uatsi. Scale = 0.6 mm. Anacroneuria varilla, new species Figs. 116-120, 140, 183-184 Adult habitus.—Head yellow except for lappets and diffuse quadrangular brown spot over ocelli. Pronotum with irregular midlateral dark bands and a pale mesal band (Fig. 116). Wing membrane transpar- ent, veins brown. Male.—Forewing length 9-10 mm. Hammer reduced to a low membranous mound or short thimble (Fig. 117). Aedea- gal apex a simple slender scoop with a pair of large ventral membranous processes; in- conspicuous lateral notches occur at base of tip. Hooks very slender, dorsal keel incon- spicuous (Figs. 118-120, 183-184). Female (putative).—Forewing length 13 mm. Subgenital plate bilobed, notch shal- 596 (A IN Figs. 151-152. A. maritza nymphal structures. 151. (151), 0.3 mm (152). low. Transverse sclerite of sternum nine in- distinct, mesal sclerite with sparse scattered setae (Fig. 140). Nymph.—Unknown. Etymology.—The species name, Span- ish for small rod, refers to the thin aedea- gal hooks and is used as a noun in appo- sition. Types.—Holotype 6 (USNM) and 6 ¢ paratypes (UMSP) from Costa Rica, Guan- acaste, Parque Nacional Guanacaste, Mar- itza, Rio Tempisquito, 550 m, 19 Jul 1987, R. Holzenthal, J. Morse, P. Clausen. Addi- tional paratypes: Costa Rica: Guanacaste: Parque Nacional Guanacaste, Rio Orosi, 700 m, 22 May 1990, R. Holzenthal, R. Blahnik, 1 3d (BPS). Parque Nacional Guanacaste, Maritza, Rio Tempisquito, 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 152 Head and pronotum. 152. Fore femur. Scales: 0.6 mm m, 17 Jun 1988, C. M. Flint, O. S. Flint, R. Holzenthal, 3 ¢6 (USNM). Estacion Marit- za, Rio Tempisquito, 8 Feb 1989, 2 6 (SWRC). Same location, 17 Mar 1989, 3 @ (INBIO). Same location, 27 Apr 1989, 3 3, 2 2 (BPS, INBIO, SWRC). Same location, 22 Oct 1990, 1 56 (SWRC). Same location, 11 May 1990, 12 3d (SWRC, INBIO). Pan- ama: Chiriqui Province: Cuenca Fortuna, Quebrada Arena 3500’, 23 May 1985, R. W. Flowers, 1 ¢ (BPS). Diagnosis.—The hammer shape and gen- eral coloration of this species are similar to A. perplexa, however the aedeagal hooks of that species are stout (Fig. 90) and the apex is trilobed (Figs. 89—90). These features eas- ily distinguish A. varilla which has slender hooks and a simple apex (Figs. 119-120). VOLUME 111, NUMBER 3 153 597 Figs. 153-154. A. uatsi nymphal structures. 153. Head and pronotum. 154. Fore femur. Scales: 0.6 mm (153), 0.3 mm (154). Anacroneuria ventana, new species Figs. 121-125 Adult habitus.—Dark Y-shaped spot cov- ers ocelli and extends to M-line; lappets dark brown. Broad median pronotal band pale, irregular midlateral bands brown, mar- ginal areas pale (Fig. 121). Wing membrane brown except for transparent window be- yond cord, veins brown. Male.—Forewing length 15 mm. Ham- mer thimble shaped, height less than basal diameter (Fig. 122). Aedeagal apex simple, tapered to a small rounded tip with a mem- branous ventral process. Hooks slender, dorsal keel moderately developed (Figs. 123-125). Female.—Unknown. Nymph.—Unknown. Etymology.—The species name, Spanish for window, refers to the transparent apical area in the wing membrane and is used as a noun in apposition. Types.—Holotype 6 from Costa Rica, Alaquela, Rio Peje, ca. 1 km SE San Vi- cente, 1450 m, 14 Feb 1992, R. Holzenthal, EF Munoz, K. Kjer (USNM). Paratype: Cos- ta Rica: Cartago: Reserva Tapanti, Quebra- da Palmitos, 1400 m, 24 Mar 1991, R. Hol- zenthal, EF Munoz, J. Huisman 1 ¢ (UMSP). San Jose: Parque Nacional Carril- lo, Rio Zurqui, 1650 m, 5 Feb 1986, J. Morse, W. Fasth, 1 ¢ (BPS). Diagnosis.—See under A. marca. Anacroneuria zapata, new species Figs. 126-130 Adult habitus.—Diffuse V-shaped pattern extends from ocelli forward to M-line; lap- pets and anterior margin of head pale brown. Pronotum brown except for narrow 598 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 155-160. Anacroneuria aedeagal structures. 155. A. magnirufa sensilla patch between hooks. 156. A. benedetto sensilla and microtrichia patch between hooks. The ridges above the patch are on the adjacent ventral sclerite. 157. A. plutonis ventral. 158. A. curiosa dorsal. 159. A. acutipennis apex, lateral. 160. A. acutipennis apex, ventral. mesal stripe and anterolateral margins (Fig. dorsal keel projects apically along mesal 126). Wing membrane transparent, veins lobe; lateral lobes of apex ear shaped (Figs. brown except for pale C and Sc. 128-130). Male.—Forewing length 13-14 mm. Female.—Unknown. Hammer cylindrical (Fig. 127). Ventral ae- Nymph.—Unknown. deagal apex trilobed, hooks slender. Small Etymology.—Zapata, Spanish for shoe, VOLUME 111, NUMBER 3 599 Figs. 161-166. Anacroneuria aedeagal structures. 161. A. annulipalpis apex, lateral. 162. A. annulipalpis apex, ventral. 163. A. benedettoi apex, lateral. 164. A. benedettoi apex, dorsal. 165. A. curiosa apex, dorsal. 166. A. curiosa apex, ventral. refers to the slipper shaped lateral aspect of Costa Rica: Alajyuela: Reserva Forestal San the aedeagal apex and is used as anoun in Ramon, Rio San Lorencito, 1090 m, 2-4 apposition. Jul 1986, R. Holzenthal, M. Heyn, B. Ar- Types.—Holotype 6 from Costa Rica, mitage, 1 ¢ (UMSP). Same location, 24— Alajuela, Rio Bochinche, Cerro Campana, 27 Feb 1987, I. Chacon, A. Chacon, 1 6 600 m, 22 Jul 1987, R. Holzenthal, J. (USNM). Guanacaste: Estacion Maritza, Morse, P. Clausen (USNM). Paratypes: Rio Tempisquito, 14 Jan 1990, 2 6 600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 167-172. Anacroneuria aedeagal structures. 167. A. divisa apex, lateral. 168. A. divisa, apex, ventral. 169. A. hacha apex, lateral. 170. A. hacha apex, ventral. 171. A. holzenthali apex, lateral. 172. A. holzenthali apex, ventral. (SWRC). Same location, 17 Mar 1989, 1 3 Diagnosis.—See under A. marca. (SWRC). Same location, 24 May 1989, 1 3 (INBIO). Same location, 22 Oct 1990, 3 Anacroneuria zarpa, new species 36 (BPS, SWRC). Puntarenas: Rio Bella- Figs. 131-135 vista, ca. 1.5 km NW Las Alturas, 1400 m, 8 Apr 1987, R. Holzenthal, S. Hamilton, M. Adult habitus.—Head patterned with dif- Heyn, 2 d (USNM, UMSP). fuse brown over most of frons; lappets VOLUME 111, NUMBER 3 464x Figs. 173-178. 601 Anacroneuria aedeagal structures. 173. A. magnirufa apex, lateral. 174. A. magnirufa apex, ventral. 175. A. marca apex, lateral. 176. A. marca apex, ventral. 177. A. marginata apex, lateral. 178. A. marginata apex, ventral. brown. Median pronotal stripe yellow, ir- regular lateral stripes pale brown with scat- tered rugosities; marginal rim black (Fig. 131). Wing membrane transparent, veins brown. Male.—Forewing length 10 mm. Ham- mer thimble shaped, height subequal to ba- sal diameter (Fig. 132). Aedeagal apex weakly trilobed and bearing a pair of dor- solateral clawlike lobes. Ventral aspect bearing prominent membranous lobes; tip rounded, hooks slender, keel absent (Figs. 133-135). Female.—Unknown. Nymph.—Unknown. Etymology.—Zarpa, Spanish for claw, re- 602 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 168m 362x 5.68 kV Figs. 179-184. Anacroneuria aedeagal structures. 179. A. perplexa apex, lateral. 180. A. perplexa apex, dorsal. 181. A. plutonis apex, lateral. 182. A. plutonis apex, ventral. 183. A. varilla apex, lateral. 184. A. varilla apex, ventral. fers to the clawlike dorsolateral lobes of the bladelike dorsolateral aedeagal lobes (Fig. aedeagal apex and is used as anoun in ap-_ 134) distinguish this species from others. position. Types.—Holotype 6 from Panama, Bo- Nomina dubia cas del Toro Province, Miramar, 21 Feb 1978, H. Wolda (USNM). Anacroneuria antica (Navas, 1924:71). Diagnosis.—The color pattern of this Type locality Costa Rica. species is not particularly distinctive but the The holotype 2 is severely damaged. Be- VOLUME 111, NUMBER 3 nedetto (pers. comm.) provided notes of an attempted artistic “reconstruction”’ of the subgenital plate. Despite this I am unable to recognize this species. Anacroneuria fulvipennis (Navas, 1934:17). Type locality San Jose, Costa Rica. The holotype @ is in fragments and de- tails of the subgenital plate are obscured (Benedetto, pers. comm.). Anacroneuria proxima Klapalek, 1923:23. Type locality Surrubres, Costa Rica. The syntypes consisting of a male and female from Costa Rica and a Mexican fe- male have not been located (Benedetto, pers. comm.). Acknowledgments I am grateful to the following individuals and institutions for the loan of specimens: R. W. Baumann, Brigham Young Univer- sity, B. C. Kondratieff, Colorado State Uni- versity, O. S. Flint and N. Adams, United States National Museum, and W. Hanson, Utah State University. I especially thank R. Holzenthal and his colleagues from the University of Minnesota and J. Jackson and D. Funk of the Stroud Water Research Cen- ter for the specimens they provided. The Stroud material was collected through sup- port of NSF Grant #BSR-9007845, and the Holzenthal material was collected through support of NSF Grants #BSR-8512368, BSR-8917684 and DEB-9400632. P. Zwick and L. Benedetto were helpful in acquiring data on type specimens and several curators and museums graciously loaned type ma- terial for study. I am also grateful to B. C. 603 Kondratieff, R. W. Baumann and R. Hol- zenthal for providing pre-publication re- views. Literature Cited Banks, N. 1914. New neuropteroid insects, native and exotic.—Proceedings of the Academy of Natu- ral Science of Philadelphia 66:608—632. Harper, P. P. 1992. Stoneflies of Panama (Plecoptera). Pp. 114-121 Jn Insects of Panama and Me- soamerica, selected studies. Oxford University Press, Oxford. Jewett, S. G. 1958. Stoneflies of the genus Anacroneu- ria from Mexico and Central America (Plecop- tera)—The American Midland Naturalist 60: 159-175. Klapalek, F 1922. Plécoptéres nouveaux.—Annales de la Sociéte Entomologique de Belgique 62:89— 95. . 1923. Plécoptéres nouveaux.—Annales de la Sociéte Entomologique de Belgique 63:21-29. Navas, R. P. L. 1924. Plecépteros, pp. 71-75. In In- sectos de la América Central—Broteria Série Zoologica 21. . 1932. Plecoptera, pp. 28-30. Jn Alcuni insetti del Museo di Zoologia della R. Universita di Torino.—Bollettino dei Musei di Zoologia e di Anatomica comparata della R. Universita di To- rino 42. . 1934. Plecdpteros, pp. 17—18. In Décadas de insectos nuevos.—Brotéria Série de Ciéncias Naturais 3. Needham, J. G., & E. Broughton. 1927. Central Amer- ican stonetlies, with descriptions of new species (Plecoptera).—Journal of the New York Ento- mological Society 35:109—120. Stark, B. P. 1995. New species and records of Anacro- neuria (Klapalek) from Venezuela.—Spixiana 18:211—249. Zwick, P. 1972. Die Plecopteren Pictets und Burmeis- ters, mit Angaben tiber weitere Arten (Insec- ta).—Revue Suisse de Zoologie 78:1123—1194. . 1973. Die Plecopteren-Arten Enderleins (In- secta): Revision der Typen.—Annales Zoologici 30:471-507. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):604—612. 1998. Studies in aquatic insects XIV: Description of eight new species of Ochrotrichia Mosely (Trichoptera: Hydroptilidae), from Costa Rica Joaquin Bueno-Soria and Ralph Holzenthal (JBS) Instituto de Biologia, Apdo Postal 70-153, Mexico 04510, D. EF; (RH) Department of Entomology, University of Minnesota, 1980 Folwell Ave, St. Paul, Minnesota 55108 U.S.A. Abstract.—The species of the following microcaddisflies are described and their male genitalia are figured: Ochrotrichia membrana, O. silva, O. avis, O. quebrada, O. dulce, O. vieja, O. ramona, and O. quinealensis. These species were collected during an inventory of the Costa Rican Trichoptera fauna. The exclusively New-World Tribe Och- rotrichiini was established by Marshall (1979) for the genera Ochrotrichia (Och- rotrichia), Ochrotrichia (Metrichia) and Rhyacopsyche. Wiggins (1996), noting lar- val characteristics that differentiated the two subgenera of Ochrotriciha, raised Me- trichia to generic status, which we also ac- cept. The species of Ochrotrichia are found from southern Canada to central Brazil and Peru, including all of the larger islands of the West Indies. Ross (1944), Denning & Blickle (1972), Blickle (1979), and Morse (1993) provided major reviews and check- lists of the described North American spe- cies. During the last 25 years, many new re- cords and descriptions of new species in the genus Ochrotrichia Mosely from the Neo- tropical Region have been published (Bo- tosaneanu & Alkins-Koo 1993; Botosa- neanu 1991, 1995; Bueno-Soria & Santia- go-Fragoso 1992, 1997; Flint 1972, 1981; Harris & Moulton 1993; Holzenthal 1988; Morse 1993). We are certain, however, that many more Neotropical species of Ochro- trichia remain to be described. Other spe- cies of Ochrotrichia described previously from Costa Rica are: O. filiforma Flint, O. pacifica Flint, and O. tenanga Flint. In the present paper we describe eight new species, thereby increasing to 11 the number of species known from Costa Rica. The species described in this paper were collected during an inventory of the Tri- choptera from Costa Rica conducted by Ralph Holzenthal and associates from 1986-1992. Morphological terminology used in the descriptions follows Marshall (1979). The holotypes will be deposited in the National Museum Natural History, Smithsonian Institution (USNM), Washing- ton, D.C., and paratypes will be deposited in the collections of the University of Min- nesota, St. Paul (UMSP) and Instituto de Biologia, Universidad Nacional Aut6noma de México (IBUNAM). Ochrotrichia membrana, new species Figs. 1—2 Diagnosis.—This species seems to be re- lated to Ochrotrichia flagellata Flint, due to the shape of the simple, elongate membra- nous lobe of tergum X. However, Ochro- trichia membrana, can be distinguished from O. flagellata by the presence of 2 short hooklike processes on the left side of the apical process of tergum X. Description of adult.—Length of fore- wing 2 mm. Color in alcohol stramineous. Male genitalia: Segment IX tubular, tergum slightly depressed but not produced anter- iad. Tergum X a large, elongate membra- nous lobe, appearing as a cylindrical flat plate in dorsal view, with 2 short apical hooklike processes on the left side; in lat- eral view, tergum X ellipsoidal, apex round- VOLUME 111, NUMBER 3 605 Figs. 1-4. Male genitalia of Ochrotrichia membrana. 1, Lateral view; 2, Dorsal view. Male genitalia of Ochrotrichia silva. 3, lateral view; 4, Dorsal view. 606 ed. Inferior appendage in lateral view, elon- gate, apex rounded; with clusters of black, peglike setae apicomesally and a row of black peglike setae ventromesally, ending in a few setae at the midbasal ridge. Phallus long, basal portion wide; apical portion nar- row, apex truncate. Material.—Holotype ¢; Costa Rica. Al- ajuela: Reserva Forestal San Ramon, Rio San Lorencito and tribs. 10°12'96"N, 84°36'42"W, 30 Mar—1 Apr 1987, 980 m, Holzenthal, Hamilton, Heyn (USNM). Etymology.—The species epithet, mem- brana, is Latin for membrane, in reference to membranous aspect of the tergum X. Ochrotrichia silva, new species Figs. 3—4 Diagnosis.—Based on the simple struc- ture and shape of tergum X, O. silva ap- pears distantly related to O. quebrada, new species. However, O. silva can be distin- guished by the presence of a short, strong, hooklike process on the right side of the basal portion of tergum X, and by the pres- ence of a long spinelike process on the left side, with its apex darker and touching the base of the apical process. Description of adult.—Length of fore- wing, 2 mm. Color in alcohol stramineous. Male genitalia: Tergum IX depressed and produced anteriad. Tergum X in dorsal view with a short, strong hooklike process, situ- ated on base of right side; middorsal portion appears membranous and cylindrical, with 1 long spinelike process on left side; apical process heavily sclerotized, slightly curved in dorsal view; in lateral view, apical pro- cess strongly curved ventrad apically. In- ferior appendage in lateral view, elongate with mid portion narrower than anterior portion, apex rounded; apical and ventral margins with a row of black, peglike setae, ending in a midbasal ridge. Phallus long and threadlike. Material.—Holotype 6; Costa Rica. Al- ajuela: Reserva Forestal San Ramon Rio San Lorencito and Tribs. 10°12'96’N, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 84°36'42”"W, 30 Mar—1 Apr 1987, 980 m, Holzenthal, Hamilton, Heyn (USNM). Etymology.—The species epithet, silva, is a Latin word for forest, in reference to the type locality, Reserva Forestal San Ra- mon. The name is treated as a noun in ap- position. Ochrotrichia avis, new species Figs. 5—6 Diagnosis.—Because of the shape of the inferior appendages in lateral view and the presence of a long spinelike process on the right side of the tergum X, O. avis is related to O. quebrada, new species. However, O. avis can be distinguished from that species by tergum X, which shows differences in the shape of the apical process, especially when viewed dorsally. Description of adult.—Length of fore- wing 2 mm. Color in alcohol stramineous. Male genitalia: Sternum, IX in lateral view, produced anteriad; tergum IX depressed and produced anteriad. Tergum X in dorsal view with elongate middorsal spinelike pro- cess on right side, apical process bifurcated apically into 2 short hooklike processes; in lateral view, on right side, apical process appears hooklike with apex ventrally di- rected and elongate middorsal spinelike process almost touching apex of hooklike process; on left side, apical process appears elongate and ventrally curved, with a short preapical process giving appearance of a bird’s neck. Inferior appendage in lateral view long, slender, with apex rounded; black peglike setae on apex and ventral margin. Phallus long with basal portion wider, at midlength surrounded and covered by a membranous structure; apical portion long and thin. Material.—Holotype 3; Costa Rica. Al- ajuela: Reserva Forestal San Ramon, Rio San Lorencito and Tribs. 10°12'96’N, 84°36'42”"W, 30 Mar—1 Apr 1987, 980 m, Holzenthal, Hamilton, Heyn (USNM). Paratype ¢, same data as holotype (UMSP). Etymology.—tThe species epithet, avis, is VOLUME 111, NUMBER 3 607 Figs. 5-8. Male genitalia of Ochrotrichia avis. 5, Right lateral view; 6, Dorsal view. Male genitalia of Ochrotrichia quebrada. 7, Right lateral view; 8, Dorsal view. derived from the Latin avis meaning bird in Ochrotrichia quebrada, new species allusion to the bird’s neck shape of the api- Figs. 7-8 cal process of tergum X. The name is treat- ed as a noun in apposition. Diagnosis.—On the basis of the presence 608 of an elongate spinelike process on the right side of tergum X and by the elongate in- ferior appendages, O. quebrada appears re- lated to O. avis. However, O. quebrada can easily be distinguished from that species by the presence of a long, dextrally curved, preapical hooklike process on tergum X. Description of adult.—Length of fore- wing 2 mm. Color in alcohol stramineous. Male genitalia: Tergum IX deeply de- pressed and produced anteriad; in lateral view, sternum IX ventrolaterally produced. Tergum X in dorsal view with long pointed process on right side; on left side a broad elongate process with apex acute and also a long dextrally curved preapical hooklike process. Inferior appendage elongate, apex rounded, with a row of black peglike setae around apex, along midventral margin and midbasal ridge. Phallus long, with basal portion wider than apical portion. Material.—Holotype ¢; Costa Rica. Guanacaste: P. N. Rincon de la Vieja Que- brada Zopilote, 10°45’9"N, 83°18'54’W, 3 Mar 1986, 785 m, Holzenthal & Fasth (USNM). Etymology.—The species epithet, que- brada, alludes to the type locality, Rincén de la Vieja quebrada Zopilote. The name is treated as a noun in apposition. Ochrotrichia dulce, new species Figs. 9-10 Diagnosis.—Based on the presence of the long process on the right side and a hooklike apical process of tergum X, O. dulce seems related to O. tenanga (Mose- ly). However, O. dulce can be distinguished from that species by the rectangular shape of the inferior appendage in lateral view with its angled anterodorsal margin. Description of adult.—Length of fore- wing 2 mm. Color in alcohol stramineous. Male genitalia: Segment IX deeply de- pressed dorsally and produced anteriad. Tergum X in dorsal view with long, straight, rodlike process on right side; body of X with long, membranous process cov- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ering most of length of an apical process that is hooked apically; in lateral view, on left side, a dorsalmost process appears, wid- er, and apically curved, ventral process ap- pears long, thinner, with its apex slightly curved upward. Inferior appendage in lat- eral view, anterodorsally angled, apical por- tion rounded, with several rows of black peglike setae; in dorsal view apex with sev- eral peglike setae and midbasal ridge with 3 black peglike setae. Phallus long, thread- like. Material.—Holotype ¢; Costa Rica. Guanacaste: Rio Tizate, 7.2 km N. E. Cafias Dulces, 10°43’98"N, 66°26'94’"W, 28 Jun 1986, 275 m, Holzenthal, Heyn, Armitage (USNM). Etymology.—The species epithet, dulce, is from the Spanish dulce, in allusion to the type locality (Canas Dulces). The name is treated as a noun in apposition. Ochrotrichia vieja, new species Figs. 11-12 Diagnosis.—On the basis of the trian- gular shape of the inferior appendages in lateral view, O. vieja appears closely related to O. cruces Flint from Mexico. However, O. vieja can be distinguished from that spe- cies by the shape and presence of 1 long process on tergum X. Description of adult.—Length of fore- wing 3 mm. Color in alcohol stramineous. Male genitalia: Tergum IX depressed and produced anteriad. Tergum X in dorsal view with first % of tergum smooth, long, apical portion rectangular; posterior % ending in a sinuous apical process; basal section of this portion with long, black tipped dorsal pro- cess, slightly curving to right apically, basal to this process is a broad thin plate mid- dorsally, with a short black-tipped spine; in lateral view posterior third of sinuous pro- cess appears wide and rounded apically and basal section of this portion bears long, black-tipped, dorsal process; basally to this process a short black-tipped spine, and a very short spur on right and left sides at VOLUME 111, NUMBER 3 609 Figs. 9-12. Male genitalia of Ochrotrichia dulce. 9, Right lateral view; 10, Dorsal view. Male genitalia of Ochrotrichia vieja. 11, Right lateral view; 12, Dorsal view. same level. Inferior appendage broadly tri- angular, apex rounded; with a band of black peglike setae apically and on midbasal ridge. Phallus slender, threadlike. Material.—Holotype 6; Costa Rica. Al- ajuela 10°46'14”N, 85°16’86’W, P. N. Rin- con de la Vieja, Quebrada Provisié6n, 4 Mar 1896, 810 m, Holzenthal and Fasth (USNM). Etymology.—The species epithet, vieja, 610 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 13-16. Male genitalia of Ochrotrichia ramona. 13, Left lateral view; 14, Dorsal view. Male genitalia of Ochrotrichia quinealensis. 15, Right lateral view; 16, Dorsal view. is from the Spanish vieja in allusion to the Ochrotrichia ramona, new species name of the locality, Rincén de la Vieja Figs. 13-14 Provision. The name is treated as a noun in apposition. Diagnosis.—Because of the presence of VOLUME 111, NUMBER 3 2 long curved spinelike processes and 1 midbasal spine on tergum X, O. ramona seems to be related to O. palitla Flint and O. felipe Ross. However, O. ramona can be distinguished from these species by the rectangular shape of the inferior appendage in lateral aspect. Description of adult.—Length of fore- wing 2 mm. Color in alcohol stramineous. Male genitalia: Tergum IX deeply de- pressed dorsally; sternum IX with a pos- terodorsal lobe. Tergum X in dorsal aspect with a small, basodorsal spine at center and 2 long, slightly curved, spines; membra- nous process ventrally, slightly twisted, with apex clearly curved to left; in lateral aspect small basodorsal spine and 2 long sinuous spines appear straight; apical por- tion of membranous process curved ventrad apically. Inferior appendage in lateral as- pect rectangular, broadest apically with dor- sal and posterior margin rectangular; pos- terior margin with many black, peglike se- tae. Phallus well developed, simple with long apical tubule. Material.—Holotype 3; Costa Rica. Al- ajuela: Reserva Forestal San Ram6én Rio San Lorencito and trib. 10°12'96’N, 84°36'42”W, 30 Mar-1 Apr 1987, 980 m, Holzenthal, Hamilton, Heyn (USNM). Paratypes: same data as holotype, 7 6 (UMSP). Etymology.—The species epithet, ra- mona, is from Spanish ‘Ramona’, feminine form of ‘Ramon’ in allusion to the name of the type locality, Reserva Forestal San Ra- mon. The name is treated as a noun in ap- position. Ochrotrichia quinealensis, new species Figs. 15-16 Diagnosis.—Because of the presence of the spines on tergum X and the elongate and broad appearance of the inferior ap- pendages in lateral view, O. quinealensis is related to O. palitla Flint. However, O. qui- nealensis can be easily distinguished by the presence, in dorsal view, of a heavy, dark, 611 spinelike process on tergum X, a character not observed in O. palitla. Description of adult.—Length of fore- wing 2 mm. Color in alcohol stramineous. Male genitalia: Sternum IX in lateral view with dorsolateral lobe; tergum IX, de- pressed and produced anteriad. Tergum X in dorsal view a broad, flat plate; at mid- length with a heavy, basally-wide spinelike process, its apex dark and directed to right; on right side, a long, slender process ending before membranous apical section, with a dark apex curved to left; apical section with a short, slender, twisted spinelike process, with dark apex curved to right; in lateral view on right side, heavy, basally-wide, spi- nelike process has its dark apex curved up- ward; twisted spinelike process of apical section appears strongly curved ventrad and partially covered by a membranous hood- like structure; on left side, heavy and wide spinelike process appears with its apex slightly curved to right, long and slender process looks like a wide plate with dark apex situated before apical section of ter- gum X, short, twisted, spinelike process of apical section seems to emerge from a hoodlike structure and is strongly curved ventrad; apical process ending in a spoon- like lobe. Inferior appendage in lateral view, elongate, broad, apex rounded, with an apical band of black peglike setae and 2, black peglike setae on midbasal ridge. Phallus long, threadlike. Material.—Holotype ¢; Costa Rica. Puntarenas; Rio Guineal, Ca. 1 km (air) E. Finca Helechales, 9°4’56"N, 83°5’52”"W, 22 Feb 1986, 840 m, Holzenthal, Morse, Fasth, (USNM). Paratypes 2d; Alajuela: Reserva Forestal San Ramon, Rio San Lorencito and tribs. 10°12'96"N, 84°36'42”"W, 30 Mar-—1 Apr 1987, 980 m, Holzenthal, Hamilton, Heyn (UMSP). Etymology.—tThe species epithet, guine- alensis, refers to the Rio Guineal, the type locality. Acknowledgments We are indebted to Dr. Oliver S. Flint, Jr., Curator of Neuropteroids of the Depart- 612 ment of Entomology of the National Mu- seum of Natural History, Smithsonian In- stitution, and to Dr. Roger Blahnik for their comments and suggestions to the manu- script; to Miss Nancy Adams, museum spe- cialist, research staff, for her assistance while we were working in the Smithsonian Institution, Washington D.C.; to Biol. Ra- fael Barba-Alvarez, for his assistance in preparing the material for study and finally to the anonymous reviewers, for their time and effort to improve the manuscript. Literature Cited Blickle, R. L. 1979. Hydroptilidae (Trichoptera) of America north of Mexico.—New Hampshire Agricultural Experimental Station Bulletin 509: 1-97. Botosaneanu, L. 1991. Amsterdam expeditions to the West Indian islands, Report 71. Trichopteres d’Haiti.—Bulletin de |’ Institut royal des Si- ences naturelles de Belgique, Entomologie 61: 113-134. . 1995. Caddis flies (Trichoptera) from the Do- minican Republic (West Indies). I. The Hydrop- tilidae.—Bulletin de Il’ Institut royal des Siences naturelles de Belgique, Entomologie 65:21—33. , & M. Alkins-Koo. 1993. Caddis flies (Insecta: Trichoptera) of Trinidad and Tobago, West In- dies.—Bulletin de |’ Institut royal des Siences naturelles de Belgique, Entomologie 63:5—45. Bueno-Soria, J., & S. Santiago-Fragoso. 1992. Studies in aquatic insects, XI: Seven new species of the genus Ochrotrichia (Ochrotrichia) from South America (Trichoptera: Hydroptilidae).—Pro- ceedings Entomological Society of Washington 94:439—446. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON , & . 1997. Studies in aquatic insects, XII: Descriptions of nineteen new species of the genus Ochrotrichia Mosely (Trichoptera: Hy- droptilidae) from Mexico and Central Ameri- ca.—Proceedings Entomological Society of Washington 99:359—373. Denning, D. G., & R. L. Blickle. 1972. A review of the genus Ochrotrichia (Trichoptera: Hydroptil- idae)—Annals of the Entomological Society of America Vol. 65:141—151. Flint, O. S., Jr. 1972. Studies of Neotropical caddis- flies, XIII: the genus Ochrotrichia from Mexico and Central America (Trichoptera: Hydroptili- dae).—Smithsonian Contributions to Zoology 118:1—28. 1981. Studies of Neotropical caddisflies, XXVIII: The Trichoptera of the Rio Limon Ba- sin, Venezuela.—Smithsonian Contribution to Zoology 330:1-61. Harris, S. C., & S. R. Moulton. 1993. New species of Ochrotrichia (Ochrotrichia) from the south- western United States and northern México. (Trichoptera: Hydroptilidae)—Journal of New York Entomological Society 101:542—549. Holzenthal, R. 1988. Catalogo sistematico de los tri- copteros de Costa Rica (Insecta:Trichoptera).— Brenesia 29:51—82. Marshall, J. E. 1979. A review of the genera of the Hydroptilidae (Trichoptera).—Bulletin of the British Museum (Natural History) 39(3):1—239. Morse, J. C. 1993. A checklist of the Trichoptera of North America, including Greenland and Mex- ico.—Transactions of the American Entomolog- ical Society 119:47—93. Ross, H. H. 1944. The caddis-flies, or Trichoptera, of Illinois.—Illinois Natural History Survey, Bul- letin 23(1):1—326. Wigginns, G. B. 1996. Larvae of the North American caddisfly genera (Trichoptera), 2nd edition. Uni- versity of Toronto Press, 370 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):613—620. 1998. A new species of the genus Gastrosaccus (Crustacea: Mysidacea: Mysidae) from Oman Masaaki Murano and Anton McLachlan (MM) Institute of Environmental Ecology, METOCEAN Co. Ltd., Riemon 1334-5, Ooigawa-cho, Shida-gun, Shizuoka 421-0212, Japan; (AM) College of Agriculture and Science, Sultan Qaboos University, PO. Box 36, Al-khod 123, Oman Abstract.—A new species, Gastrosaccus trilobatus, is described based on specimens from sandy beaches of Oman. The new species is distinguished from known species of the genus by the carapace and telson. The carapace is pro- vided with 3 posteriorly directed lobes on the posterodorsal margin, and the telson is armed with 2 distal conspicuously large spines and 4 or 5 spinules inserted between them. The genus Gastrosaccus was established by Norman in 1868, and now contains 17 species as far as we can determine. List of these species, in company with their local- ities, are shown in Table 1. It is distin- guished from related genera by the third male pleopod with the styliform exopod and the multi-segmented endopod, the uni- ramous female pleopods except for the first pair which is biramous, and the labrum without accessory spines in both sides of the median spiniform process. The present new species was collected during an eco- logical study of sandy beaches in Oman, Arabian Peninsula, by one of the authors (AM). The type specimens are deposited in the Zoological Museum, University of Co- penhagen. Gastrosaccus trilobatus, new species Figs. 1-3 Type series.—Holotype (CRU 3411), adult male (6.0 mm); allotype (CRU 3412), gravid female (7.3 mm); paratypes (CRU 2419), 1 adult gravid female (5.8 mm), 3 immature males (4.6, 3.5, 3.3 mm) and 1 immature female (3.5 mm); Khaluf, Oman, 20°25’N, 58°00’E, sandy beach, 6 Novem- ber 1995, coll. A. McLachlan. Other para- types (CRU 2420), 2 adult males (5.6, 6.5 mm), 1 adult female (5.5 mm) and 1 im- mature female (5.4 mm), Majis, Oman, 24°30'N, 56°40’E, sandy beach, 27 Septem- ber 1995, coll. A. McLachlan; 2 specimens of which, male of 6.5 mm and adult female, were dissected for drawing. Description.—Rostrum low triangular with narrowly rounded apex, somewhat bent downwards, covering basal part of eyestalks (Fig. 1A, B). Anterolateral corner of carapace rounded. Posterior margin of carapace deeply emarginate, leaving last thoracic somite uncovered dorsally, split in each side of emargination, overlapping with posterolateral margin; emarginated part fur- nished with 3 lobes, median lobe elongate triangular with pointed apex, side lobes bill- shaped, directed posteriorly, not reflected; posterior and posterolateral margins from side lobe to posterolateral angle weakly ser- rated (Fig. 1C). Posterolateral angle of car- apace rounded, covering side of first ab- dominal somite. Eye well developed; cornea as wide as or slightly wider than stalk (Fig. 1A, B). Antennular peduncle of male more robust than that of female; first segment shorter than combined length of second and third segments, with 4 short setae at anterolateral 614 Table 1.—List and localities of 18 Gastrosaccus species. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Species . australis W. M. Tattersall . bispinosa Wooldridge . daviei Bacescu & Udrescu . dunckeri Zimmer QAAQA Q . kempi W. M. Tattersall . longifissura Wooldridge . Mediterraneus Bacescu . msangi Bacescu ae) @ Lachlan olivae Bacescu . robusta Panampunnayil . roscoffensis Bacescu . sanctus (van Beneden) AARAAD G. sorrentoensis Wooldridge & Mc- Lachlan G. spinifer (Goés) G. trilobatus, new species . brevifissura O. S. Tattersall namibensis Wooldridge & Mc- psammodytes O. S. Tattersall Localities New Zealand S. Africa S. Africa NE Australia Between Ceylon and New Guinea India E. Africa Singapore waters S. China Sea India S. Africa Arabian Gulf S. Africa Mediterranean Tanzania Namibia Orange River estuary S. Africa SE Australia France Atlantic coasts of Europe and Africa, Mediterranean W. Australia Atlantic coasts of Europe and Africa, Mediterranean Oman Authorities W. M. Tattersall 1923 Wooldridge 1978 O. S. Tattersall 1952, 1962 Bacescu & Udrescu 1982 Zimmer 1915 W. M. Tattersall 1922, Pillai 1973 O. S. Tattersall 1958 O. S. Tattersall 1960 li 1964 W. M. Tattersall 1922 O. S. Tattersall 1962 (as G. gordonae) Murano (1998) Wooldridge 1978 Bacescu 1970 Bacescu 1975 Wooldridge & McLachlan 1987 O. S. Tattersall 1955 (as G. sanctus) O. S. Tattersall 1958 Panampunnayil 1989 Bacescu 1970, Nouvel 1972 Tattersall & Tattersall 1951 Wooldridge & McLachlan 1986 Tattersall & Tattersall 1951, Lagardére & Nouvel 1980 Present record corner; second segment shortest, wider than long, with 3 spines on dorsolateral surface and 1 short and 1 long setae at inner distal corner; third segment 1.5 times longer than wide, with 2 spinules on dorsal surface, no- ticeable digitate process present on dorsal surface near base of outer flagellum. Fe- male antennular peduncle with first segment as long as second and third segments com- bined, second segment with 2 long and 1 short setae at outer distal corner. Outer fla- gellum with basal lobe fringed with sensory setae (Fig. 1A, B). Antennal scale extending to distal end of second segment of antennular peduncle in male, slightly less in female, slightly more than 3 times longer than broad; lateral mar- gin slightly convex, naked, terminating in spine extending to apex of scale, distal su- ture distinct (Fig. 1D). Antennal peduncle shorter than antennular peduncle, longer than antennal scale; second segment lon- gest, more than twice longer than broad, armed on inner margin with 4 plumose se- tae of which distal one is longest and thick- est; third segment 0.4 as long as second, with 3 plumose setae on inner margin of which distal one is longest and thickest (Fig. 1D). Labrum longer than broad, with long me- dian spiniform process (Fig. 1H). Mandib- ular palp slender, third segment 7% of second segment in length (Fig. 1E). Maxillule and maxilla as shown in Fig. 1F and G, respec- tively. Hook-shaped anteromedian process present on ventral side of clypeus (Fig. 11). First and second thoracic limbs with en- dopod rather slender (Fig. 2A, B). Endo- pods of third to eighth limbs similar in shape, ischium longer than merus, ischium VOLUME 111, NUMBER 3 615 Fig. 1. Gastrosaccus trilobatus, new species, A, D, E, G, I, male (6.5 mm); B, C, EK H, female (5.5 mm). A, anterior end in dorsal view; B, anterior end in dorsal view; C, posterodorsal margin of carapace; D, antenna; E, mandible and mandibular palp; E maxillule; G, maxilla; H, anterior end in ventral view; I, anteromedian process of clypeus (APC) and anterior part of labrum (LA) in lateral view. 616 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.5 ee 0.2mm A Fig. 2. Gastrosaccus trilobatus, new species, male (6.5 mm). A, endopod of first thoracic limb; B, endopod and proximal part of exopod of second thoracic limb; C, endopod of third thoracic limb; D, fifth thoracic limb; E, seventh thoracic limb. VOLUME 111, NUMBER 3 and merus shortened and carpopropodus lengthened toward posterior pairs, carpo- propodus subdivided into 6 to 10 subseg- ments increasing in number toward poste- rior pairs (Fig. 2C—E). Basal plate of tho- racic exopods with distolateral corner point- ed in second to seventh limbs (Fig. 2B, D, E) and rounded in first and eighth limbs. Flagella of thoracic exopods 10-segmented in first limb, 11 in second and third, 12 in fourth to eighth (Fig. 2D, E). Abdomen with second and third somites shortest, fifth somite 1.5 times longer than fourth, sixth somite 1.2 times longer than fifth. All pleopods of male biramous. First pleopod with 8-segmented exopod and un- segmented endopod, sympod with 9 plu- mose setae along lateral margin (Fig. 3A). Second pleopod with 7-segmented endopod and 8-segmented exopod, exopod much broader and longer than endopod, distal 3 segments extending beyond apex of endo- pod, each of proximal 4 segments armed at outer distal corner with thick seta, proximal part of which has undulate outer margin (Fig. 3B). Third pleopod: exopod extremely elongate, styliform, exceeding distal end of sixth abdominal somite, 4-segmented, first segment longest, indistinctly divided into about 4 subsegments, second segment. as long as third and fourth segments together, second and third segments unarmed, fourth (distal) segment shortest, 0.37 as long as third, armed with 2 strong, subequal, barbed setae on distal end and short seta on distal third of lateral margin; endopod 6-seg- mented, extending beyond middle of first segment of exopod (Fig. 3C). Fourth pleo- pod with unsegmented endopod and 8-seg- mented exopod (Fig. 3D). Fifth pleopod al- lied to fourth but slightly smaller, endopod unsegmented, exopod 7-segmented (Fig. 3E). Female pleopods: first pleopod bira- mous, endopod unsegmented, shorter than exopod, with naked seta at tip; exopod un- segmented, with 3 plumose setae on distal end; sympod cylindrical, outer margin with 2 long setae near proximal end and 3 setae 617 on distal end (Fig. 3F). Second to fifth ple- opods uniramous, slender (Fig. 3G). Endopod of uropod longer than exopod, with 8—10 strong spines on ventral inner margin from statocyst region to near distal end. Exopod of uropod truncate distally, as long as telson, armed along lateral margin with 12 strong spines arranged regularly (Fig. 31). Telson slightly longer than last abdomi- nal somite, 2.3 times as long as maximum width at base; cleft more than % of telson length, armed with 15—20 spinules on either side; each apex of distal lobes with 1| strong spine; lateral margin nearly straight, with 6—7 large spines, distal one conspicuously larger than others and slightly larger than apical spine; 5 and 1 spinules inserted be- tween apical spine and distal conspicuously large lateral spine and between fourth and fifth large spines, respectively (Fig. 3H); single spine present on median anteroven- tral line. Etymology.—Derived from 3 lobes on the posterodorsal margin of the carapace. Remarks.—In the new species the telson bears spinules between the apical spine and the distal spine of the lateral margin. To date, such a telson is recorded in 5 species of the genus Gastrosaccus, G. bispinosa Wooldridge, 1978, G. brevifissura O. S. Tattersall, 1952, G. kempi W. M. Tattersall, 1922, G. longifissura Wooldridge, 1978, and G. msangi Bacescu, 1975. The new species is easily distinguished from them by the 3 lobes on the posterodorsal margin of the carapace and the 2 conspicuously large distal spines on the telson. There are intraspecific variations in the depth of the apical cleft and the armature on the lateral margin of the telson. In an adult female (5.8 mm) from Khaluf, the api- cal cleft of the telson is shallower (about of telson length), more divergent posteri- orly and the spinules are fewer in number. Moreover, the lateral margin of the telson is armed with 7 large spines, and the num- ber of inserted spinules is one each in spac- es between fourth and fifth, fifth and sixth, 618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON TT we Ee Fig. 3. Gastrosaccus trilobatus, new species, A—E, H, I, male (6.5 mm); KE G, female (5.5 mm); J, female (5.8 mm). A, first pleopod; B, second pleopod; C, third pleopod; D, fourth pleopod; E, fifth pleopod; F first pleopod; G, third pleopod; H, telson; I, uropod in ventral view; J, telson. VOLUME 111, NUMBER 3 and sixth and seventh large spines and 4 in the space between the seventh and apical spines (Fig. 3J). A prominent spine on the median line of the anteroventral surface of the telson was found in the new species. A similar spine was reported for Gastrosaccus sorrentoen- sis and G. psammodytes. As noted by Wooldridge & McLachlan (1986), it may be a common character in the genus Gastro- saccus. Ecological note.—Gastrosaccus triloba- tus was found on all 10 beaches surveyed in Oman in numbers ranging from 54—4128 per meter transect and up to 300 individu- als/m?. It tended to be more common on the most exposed beaches. Intertidal distribu- tion of the mysids during low tide was across the saturated lower shore and into the surf zone. Numbers peaked between the mid tide and spring low tide levels. The beaches in Oman have fine sand and are mesotidal with a maximum tide range of 3 m. Wave energy is low except in the south. Water temperature and salinity on the sam- pling days were 27—33°C and 31—36 PSU, respectively. More ecological information is forthcoming. Acknowledgements The first author thanks Dr. Niel L. Bruce, Zoological Museum, University of Copen- hagen, for giving him the opportunity to ex- amine the present material. Literature Cited Bacescu, M. 1970. Contributions a |’étude morpho- écologique des Gastrosaccinae (Crustacea, Mysidacea) du versant est de 1’ Atlantique et de la Mediterranée.—Revue Roumaine de Biolo- gie, Série de Zoologie 15:217—234. . 1975. Contributions to the knowledge of the mysid (Crustacea) from the Tanzanian wa- ters.—University Science Journal, University of Dar es Salaam 1:39-61. , & A. Udrescu. 1982. New contribution to the knowledge of the Mysidacea from Australia.— Travaux du Muséum d’Histoire naturelle Gri- gore Antipa 24:79—96. li, N. 1964. Fauna Japonica, Mysidae (Crustacea).— 619 Biogeographical Society of Japan, Tokyo, 610 Pp. Lagardeére, J.-P, & H. Nouvel 1980. Les Mysidacés du talus continental du golfe de Gascogne. II. Fam- illes des Lophogastridae, Eucopiidae et Mysidae (Tribu des Erythropini exceptée).—Bulletin du Muséum national d’ Histoire naturelle, Paris, 4° série, 2, section A, no 3:845—887. Murano, M. 1998. Mysidae (Crustacea: Mysidacea) collected from the western Arabian Gulf.— Plankton Biology and Ecology 45:45—54. Norman, A. M. 1868. Preliminary report on the Crus- tacea, Molluscoida, Echinodermata, and Coe- lenterata, procured by the Shetland Dredging Committee in 1867.—Report of the British As- sociation for the Advancement of Science, Lon- don 37(1867):437—441. Nouvel, H. 1972. Observations sur les Mysidacés et quelques Cumcés littoraux de la Céte fran- caise du golfe de Gascogne au sud de lV’embouchure de la Gironde.—Bulletin. Cen- tre d’Etudes et de Recherches scientifiques, Biarritz 9:127—140. Panampunnayil, S. U. 1989. A new species of Gastro- saccus (Crustacea: Mysidacea) from the South West Coast of Australia.—Journal of Plankton Research 11:1307-1314. Pillai, N. K. 1973. Mysidacea of the Indian Ocean.— Indian Ocean Biological Centre, Handbook 4: 1-125. Tattersall, O. S. 1952. Report on a small collection of Mysidacea from estuarine waters of South Af- rica.—Transactions of the Royal Society of South Africa 33:153-187. . 1955. Mysidacea.—Discovery Reports 28:1— 190. . 1958. Further notes on Mysidacea from South African waters.—Transactions of the Royal So- ciety of South Africa 35:373—385. . 1960. Report on a small collection of Mys- idacea from Singapore waters.—Proceedings of the Zoological Society of London 135: 165-181. . 1962. Report on a collection of Mysidacea from South African off-shore and coastal waters (1957-59) and from Zanzibar (1961).—Pro- ceedings of the Zoological Society of London 139:221—247. Tattersall, W. M. 1922. Indian Mysidacea.—Records of the Indian Museum 24:445—504. . 1923. Crustacea. Part 7. Mysidacea.—British Antarctic (“Terra Nova’) Expedition, 1910, Natural History Report, Zoology 3:273-304. , & O. S. Tattersall 1951. The British Mysida- cea. The Ray Society, London, 460 pp. Wooldridge, T. 1978. Two new species of Gastrosac- cus (Crustacea, Mysidacea) from sandy beaches 620 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in Transkei.—Annals of the South African Mu- saccus (Mysidacea) from the south—west coast seum 76:309-327. of Africa.—Crustaceana 52:101—108. , & A. McLachlan, 1986. A new species of Zimmer, C. 1915. Schizopoden des Hamburger Natur- Gastrosaccus (Mysidacea) from Western Aus- historischen (Zoologischen) Museums.—Miittei- tralia——Records of the Western Australian Mu- lungen aus dem Naturhistorischen (Zoologisch- seum 13:129—-138. en) Museum 32:159—-182. , & . 1987. A new species of Gastro- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):621—626. 1998. A new species of amphipod (Crustacea: Amphipoda: Lysianassoidea) from the Pacific Coast of North America Ann Dalkey Environmental Monitoring Division, City of Los Angeles, 12000 Vista Del Mar, Playa del Rey California 90293, U.S.A. Abstract.—A new species of lysianassoid amphipod, Lepidepecreum serra- culum, is described. This species possesses a distinctive rudder-shaped convex carination of urosomite | that combined with a slightly carinate antenna 1, chelate gnathopod 2, and rounded corners of the epimera, distinguishes it from other species of Lepidepecreum. It occurs in a variety of substrata subtidally to 150 m depth from the Mexican—American border to Alaska. An undescribed species of the lysianas- soid amphipod Lepidepecreum has been en- countered regularly in sublittoral surveys conducted in southern California during the past 20 years. This species was first rec- ognized by Barnard (1969) who compared it with several Lepidepecreum species in- cluding Lepidepecreum gurjanovae Hurley, 1963. Barnard, noting that his specimens differed morphologically from L. gurjano- vae and occurred in much shallower water, listed his material as Lepidepecreum ? gur- jJanovae pending further study. Subsequent- ly, during Southern California Association of Marine Invertebrate Taxonomists (SCAMIT) workshops conducted with the late J. Laurens Barnard, the species was recognized as distinct, both morphological- ly and ecologically. It was listed as Lepi- depecreum sp. A prior to formal description (SCAMIT 1996). Lepidepecreum serraculum, new species Figs. 1-3 Material examined.—All specimens have been deposited in the Los Angeles County Museum of Natural History. Holotype: male, Santa Monica Bay, City of Los An- geles, Environmental Monitoring Division, station DN8, 33°51.02'N, 118°25.00'W, 7 Feb 1986, 22 m depth, LACMNH No 86- 523.1. Paratypes: Huntington Beach, Cali- fornia, County Sanitation Districts of Or- ange County Station B4, 17 Jul 1980, 33°34.54'N, 117°59.75'W, 60 m depth, one female, LACMNH No. 80-162.1, Station BO(1) 5 Feb 1980 33°34.67'N, 118°00.54"W 60 m depth, one female, LACMNH No 80- 163.1, and Station Control(1), 15 Jan 1981, 33°35.95’'N, 118°03.79'W, 60 m depth, one female, LACMNH No 81-240.1; Santa Monica Bay, City of Los Angeles, Environ- mental Monitoring Division Station D3, 33°51.78'N, 118°35.25'W, 24 Jan 1994, 78 m depth, one female, LACMNH No 94-90. 1 and Station E8, 33°54.30'N, 118°36.43’W, 29 Jul 1993, 151 m depth, two males, LACMNH No 93-143.1. Diagnosis.—Coxae and pereopods 5-7 ornamented with fine setules. Antenna 1, article 1 slightly carinated, callynophore present in males. Metasomite 3 with pos- terodorsal carina. Urosomite 1 with pro- nounced rudder-shaped carina. Mandible with moderately developed setulose molar. Maxilla | outer plate spines well developed. Gnathopod 2 minutely chelate. Pereopods 5—7 article 2 posterior margins moderately serrate. Epimeron 3 posterior margin straight with rounded posteroventral corner. Uropods and telson spinose. Description.—Male, body smooth with thick white integument, metasomite 3 bear- 622 ing posterodorsal carina, urosomite 1 with pronounced rudder-shaped carina (Figs. la, b). Coxae 1—4 longer than broad, slightly expanded distally (ventrally), partially cov- ered with fine setules and bordered with many setules; coxa 4 excavate posteriorly. Head subequal in length to first pereon- ite, lateral lobes of head strongly produced, elongate eyes with deep reddish-brown pig- mented ommatidea in alcohol. Antenna 1 short; peduncle robust, article 1 dorsal mar- gin carinate bearing plumose setae dorsally and on distoventral corner, article 2 half as long as article 1, article 3 one third as long as article 1; accessory flagellum with 4 ar- ticles; main flagellum longer than peduncle, article 1 with callynophore of aesthetascs as long as the accessory flagellum (Fig. Ic). Antenna 2 long, nearly as long as body length; 8 setae subapically on outer ventro- lateral margin of peduncular article 3, arti- cles 4 and 5 both possessing a row of se- tules on dorsal margin and single plumose seta on anteroventral corner; flagellum at least 60-articulate, the first 40 calceolate (Fig. 1d). Epistome broadly rounded pro- duced above upper lip. Mandible with mod- erately developed setulose molar, row of raker spines, incisor well defined with con- vex cutting edge; palp proximal to molar, 3-articulate, article 1 short, article 2 long bearing distal spines, article 3 half as long as article 2 with inner and distal spines (Fig. le). Maxilla 1: inner plate bearing distal plumose setae and many fine hair-like setae; outer plate with 11 multicuspidate spine- teeth; palp 2-articulate, article 2 longer hav- ing small distal teeth and a seta (Fig. 1f). Maxilla 2: inner plate with row of fine hair- like setae on inner proximal margin, several setae and serrated spines on inner distal margin; outer plate bearing closely bundled setae and serrate spines distally (Fig. 1g). Maxilliped: inner plate with apical stout teeth and spines on distal margin and plu- mose setae along inner margin; outer plate bearing basal setae, peg-like spines along the inner margin and small facial spines ad- jacent to peg-like spines; palp 4-articulate, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON article 2 longest with setae on inner margin and a distal seta on outer margin, article 3 similar to article 2 though shorter and bear- ing several distal setae, article 4 covered with striations or possibly short setules (not illustrated), prominent apicomedial seta on outer margin and small terminal nail (Fig. 1h). Gnathopod 1 subchelate; dactyl with proximal tooth that forms a bifid process; article 6 sub-rectangular with distal setae, palmar margin bearing 2 medial teeth, ter- minating with 2 or more spines, dactyl with subterminal tooth; article 5 subequal in length to article 6; articles 3 and 4 short, subequal with distal setae; article 2 as long as articles 3—6 combined bearing several se- tules along anterior margin and setae on distoposterior corner (Fig. 2a). Gnathopod 2 minutely chelate; dactyl small, triangular bearing a single seta; article 6 anterior mar- gin convex bearing 3 rows of serrate setae, lower margin straight, terminating in tooth armed with small spines; article 5 twice as long as article 6, anterior margin convex with many fine setules and setae distally, posterior margin expanded distally covered with a rasp of many scales; article 4 shorter than article 5, expanded distally with prom- inent anterior setae; article 3 length sub- equal to article 5; article 2 length subequal to combined length of articles 3—6 (Fig. 2b). Pereopod 3 simple; locking spines on distal end of article 6, article 4 expanded along anterior margin (Fig. 2c). Pereopod 4 similar to pereopod 3 in size and shape (Fig. 2d). Pereopod 5 shorter than pereo- pods 3 and 4; article 4 expanded distopos- teriorly, lobe reaching over nearly half of article 5; article 2 ovoid, covered with fine setules, posterior margin moderately ser- rate; coxa subquadrate, covered with fine setules and short spines (Fig. 2e). Pereopod 6 length subequal to pereopod 5; article 4 expanded distoposteriorly; article 2 ovoid with posterior margin moderately serrate (Fig. 2f). Pereopod 7 shorter than pereo- pods 5 and 6; article 4 also expanded dis- toposteriorly; article 2 subquadrate, length VOLUME 111, NUMBER 3 623 Fig. 1. Lepidepecreum serraculum, new species: a, entire holotype with antenna 2 placed outside coxal plates to illustrate relative length; b, urosome; c, antenna 1; d, antenna 2; e, mandible; f, maxilla 1; g, maxilla 2; h, maxilliped. 624 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Lepidepecreum serraculum, new species: a, gnathopod 1; b, gnathopod 2; c, pereopod 3; d, pereopod 4; e, pereopod 5; f, pereopod 6; g, pereopod 7. VOLUME 111, NUMBER 3 625 Fig. 3. subequal to articles 4—7 combined, poste- rior margin moderately serrate with small, distoposterior excavation that varies in de- gree among and within individuals (Fig. 2s) Epimera 1—2 posteroventrally convex; epimeron 3 (Fig. la) posterior margin Straight with rounded posteroventral corner. Uropod 1 long; peduncle dorsally excavate with spinose margins; spinose rami slightly shorter than peduncle, both with spines and subterminal nail (Fig. 3a). Uropod 2 shorter than uropod 1; peduncle dorsally excavate with spines and setules on margins; rami spinose and subequal in length, both with subterminal nail (Fig. 3b). Uropod 3 pedun- cle shorter than uropod 1 peduncle, deeply excavate dorsally with setae on outer face and spines on medial margin; inner ramus subequal to peduncle in length, foliaceous, bearing setules on the proximal-lateral mar- gin and spines on the apicolateral margin; outer ramus longer, foliaceous with many plumose setae on beveled apicolateral mar- gin (Fig. 3c). Telson as long as uropod 3 outer ramus, longer than broad, incised three-fourths of its length, each lobe taper- ing distally, each with small apical plumose seta, apical spines and subterminal spine and setule (Fig. 3d). Female, non-ovigerous: Like male but antenna | short, flagellum only 6-articulate, Lepidepecreum serraculum, new species: a, uropod 1; b, uropod 2; c, uropod 3; and d, telson. lacking callynophore; antenna 2 lacking calceoli; uropod 2 lacking plumose setae. Remarks.—Other species differ from L. serraculum in cuticle ornamentation, body and antenna | carination, gnathopod che- lation, pereopods 5-7 serration, uropod and telson spination. Lepidepecreum serracu- lum is most similar to L. gurjanovae which differs from the former by possessing for- ward directed carination on article 1 of the first antennal peduncle, a sub-chelate gnathopod 2, carination on all pereonites, subrectangular epimeron 3 pleon epimeron, and upturned carination of urosomite 1 (Hurley 1963). Although their geographic ranges overlap in the northeastern Pacific, the species are separated bathymetrically (Lepidepecreum serraculum subtidal to 150 m, L. gurjanovae >260 m). Another north- eastern Pacific species L. garthi differs from L. gurjanovae by the absence of eyes, forward directed carina on article 1 of the first antennal peduncle, pronounced cari- nation on all body segments, subchelate gnathopod 2, and fewer uropod and telson spines (Hurley 1963). Other northeastern Pacific species L. comatum and L. vitjazi differ from L. serraculum by possessing a coarser cuticle ornamentation; L. alectum and L. kasatka by lacking carination on an- tenna 1; L. eoum and L. rostratum by pos- sessing greater body carination; and L. nau- 626 tilus and L. umbo in differing body cari- nation and by lacking uropods and telson spines (Gurjanovae 1962, Hirayama 1985, Barnard and Karaman 1991). A freshly preserved Lepidepecreum ser- raculum specimen had orange-pink dots at the base of coxal plates 1—3 and 5; orange- pink dots on the dorsum of pereonites 1—9 including a band across pereonite 8. Etymology.—The specific epithet serra- culum is a Latin neuter noun, meaning rud- der, and is placed in apposition with the ge- neric name. It alludes to the easily recog- nizable rudder-shaped dorsal carina on uro- somite 1. Distributional ecology.—This species occupies a wide range and variety of habi- tats. It occurs from the Mexican—American border and extends north along the Pacific Coast to Alaska. Typical specimens from southern California are 3 mm in length whereas those from northern range of dis- tribution are twice as long (Norma Jarrett, pers. comm.). It is found in sediments rang- ing from fine (sandy silt) to coarse (red sands), off open ocean coast lines and in harbors. Depth range is from intertidal to 150 m. Acknowledgments Special appreciation is extended to Ms. Norma Jarrett, who helped me in my initial efforts to identify this species and provided information on the northern specimens and to the late Dr. J. Laurens Barnard, who sup- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON plied encouragement after reviewing the first written draft. Comments from Charles Oliver Coleman and three additional re- viewers resulted in significant improve- ments to this manuscript. This paper was partially funded by SCAMIT Grant Num- ber 98-1 and is SCAMIT Contribution Number 13. Literature Cited Barnard, J. L. 1969. Gammaridean amphipoda of the rocky intertidal of California: Monterey Bay to La Jolla—— Bulletin of the United States Nation- al Museum 258:1—230. , & G. S. Karaman. 1991. The families and genera of marine gammaridean Amphipoda (ex- cept marine Gammaroids).—Records of the Australian Museum, Supplement 13(Part 2): 419-866. Gurjanovae, E. E 1962. Bokoplavy severnoi chasti Tixogo Okeana (Amphipoda-Gammaridea) chast’ 1.—Akademii Nauk SSSR, Opredeliteli po Faune SSSR 74:1—440. Hirayama, A. 1985. Taxonomic studies on the shallow water Gammaridean amphipoda of West Kyu- shu, Japan. V. Leucothoidea, Liljeborgiidae, Lysianassidae (Prachynella, Aristias, Ensayara, Lepidepecreum, Hippomedon and Anonyx).— Publication of the Seto Marine Biological Lab- oratory 30:167—212. Hurley, D. E. 1963. Amphipoda of the family Lysi- anassidae from the west coast of north and cen- tral America.—Allan Hancock Foundation Pub- lications, Occasional Paper 25:1—165. Southern California Association of Marine Inverte- brate Taxonomists (SCAMIT). 1996. A taxo- nomic listing of soft bottom macro- and me- gainvertebrates from infaunal and epibenthic monitoring programs in the Southern California Bight. Edition 2, San Pedro, California, 86 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):627—633. 1998. Metatiron bonaerensis, a new species (Crustacea: Amphipoda: Synopiidae) from the southwest Atlantic Gloria M. Alonso de Pina Museo Argentino de Ciencias Naturales ‘““‘Bernardino Rivadavia,”’ Divisi6n Invertebrados, Avda. Angel Gallardo 470, (1405) Buenos Aires, Argentina Abstract.—A new species, Metatiron bonaerensis, is described and illustrated from the south of Buenos Aires province, Argentine continental shelf, south- west Atlantic. The new species is assigned to Metatiron based on the absence of mandibular palp. M. bonaerensis is separated from Tiron tropakis by the laterally smooth pleonites 1—3, protuberant forehead, quadrate coxa 7 and max- illa 1 inner plate shape. The relationship with other species of the genus is discussed. The material was dredged at different depths and the grain size of the sediment was determined for each sampling station. The new species described here was dis- covered from the El] Rinc6n area of the Ar- gentine continental shelf (Buenos Aires province), approximately 39° to 40°S and 61° to 62°W. Benthic samples were collect- ed using a Van Veen dredge during the sur- vey carried out by the R/V El Austral in 1993. The type specimens are deposited in the Museo Argentino de Ciencias Naturales ‘““Bernardino Rivadavia,’’ Buenos Aires, Argentina (MACN). Metatiron Rabindranath, 1972 Metatiron bonaerensis, new species Figs. 1-49 Holotype.—Male 6.75 mm (MACN 34004), El Rincdén, Buenos Aijres: 39°39'40"S, 61°50’01"W (sta 24); depth 15 m; 2 Nov 1993. Allotype.—Ovigerous female 9.6 mm (MACN 34005), El Rinc6n, Buenos Aires: 39°25'10"S, 61°34’57”W (sta 19); depth 18 m; 69% fine sand; 3 Nov 1993. Paratypes.—1 male ca. 5.0 mm, | juve- nile ca. 2.0 mm (MACN 34006), El Rincon, Buenos Aires: 39°05’36"S, 61°20'11”W (sta 3); depth 9.5 m; 78% medium sand; 3 Nov 1993. 1 ovigerous female 6.0 mm (MACN 34007), El Rinc6én, Buenos Aijres: 39°20'08"S, 61°25'02”W (sta 15); depth 19 m; 69% fine sand; 3 Nov 1993. | immature female ca. 4.0 mm (MACN 34008), El Rin- con, Buenos Aires: 39°30’07"S, 61°34'58”W (sta 22); depth 18 m; 75% fine sand; 3 Nov 1993. 1 female with oostegites ca. 6.0 mm, 1 immature male 6.0 mm (MACN 34009), same data as holotype. 1 ovigerous female 5.8 mm (MACN 34010), El Rincén, Bue- nos Aires; 39°40'07"S, 61°35’05”W (sta 25); depth 19 m; 67% fine sand; 2 Nov 1993. Description.—Male holotype, body length 6.75 mm. Head about as long as first 3 peraeonites combined; forehead protuber- ant, dorsal margin forming right angle an- teriorly; rostrum short, pointing acutely over base of antenna 1; lateral cephalic lobe moderately produced. Eyes well-developed, of medium size; accessory eye composed of 2 separated ommatidia (Fig. 1). Antenna 1 somewhat shorter than peduncle of antenna 2; peduncle article 1 broad, longer than pe- duncle articles 2 and 3 combined, with sin- gle distal spine anteriorly, which is as long as peduncle article 2; peduncle article 2 about one-third as long as peduncle article 1; peduncle article 3 shorter than article 2, almost one-half the length of this latter; fla- 628 gellum with 9 articles, article 1 elongate about one-third the length of flagellum, bearing fringe of setae; accessory flagellum with 6 articles, slightly longer than flagel- lum articles 1 and 2 combined (Fig. 2). An- tenna 2 elongate, about as long as the body; peduncle articles with fine setae anteriorly; peduncle articles 4 and 5 very long, sub- equal in length; flagellum with 17 articles (Fig. 3). Upper lip and epistome as figured (Fig. 4). Mandible without palp; incisor with 4 teeth on right mandible and 2 more accessory teeth on the left; molar promi- nent, columnar, with triturating surface and plumose setae; lacinia mobilis large and dentate; spine row consisting of 6 spines subequally elongate on right mandible and 7 spines on the left mandible, being one of them shorter than the other ones (Figs. 5— 7). Lower lip with well-developed mandib- ular lobes, inner lobes present and separate from each other (Fig. 8). Maxilla 1, inner plate fully setose, bearing 3 setae at the apex separated from the rest of medial setae by a smooth acclivity (Fig. 9); outer plate with 7 apical spines (Fig. 10); palp carrying 7 terminal tooth-spines and 4 subterminal setae (Fig. 11). Maxilla 2, inner plate broader than outer with dense medial and submarginal setal row (Fig. 12). Maxilliped as illustrated (Fig. 13). Gnathopods 1 and 2 similar, simple; propodus thin, about 0.5 length of carpus, posterior margin with pectinate spines; dac- tylus bearing inner tooth. Gnathopod 1, coxa distally expanded; basis shorter than the next 3 articles combined, slightly broad- er medially, with long plumose setae along anterior margin; carpus elongate, posterior margin with pectinate spines (Figs. 14, 15). Gnathopod 2, coxa narrow; basis about as long as the next 3 articles combined, almost straight, with long plumose setae on distal half of anterior margin and along posterior margin; carpus elongate, posterior margin armed with spines and long plumose setae (Figs. 16, 17). Peraeopods 3—7, propodus short and broad; dactylus stubby and bear- ing an inner tooth. Peraeopods 3 and 4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON small (Figs. 18—21), similar except for coxa; coxa of peraeopod 3 large, distally expanded and posteriorly with a lobe; coxa of peraeopod 4 small, short and broad; dac- tylus of both appendages imbeded in end of propodus, forming a subspherical disk with hooked tooth, a long seta matching curve of hook, and an accessory small spine at base of hook. Peraeopods 5 and 6 similar; coxa with posterior lobe; basis ovate, longer than wide, with distal posterior lobe (Figs. 22-25). Peraeopod 7, coxa nearly oval; ba- sis broader than long, almost quadrate, weakly crenellate posteriorly; merus broad- er than on peraeopods 5 and 6 (Figs. 26, 27). Peraeopods 5—7, dactylus not spherical basally and only hooked distally. Pleonal epimera 1—3, posteroventral cor- ners slightly produced, posterior margins bearing some notches with setules (Figs. 28-30). Pleonites 1—3 crenellate posteriorly on dorsal edge, crenellations superimposed, almost indistinguishable, small and round- ed; lateral segmental margins evenly smooth (Fig. 31). Urosomites 1 and 2 with medium and large dorsal teeth, respectively; urosomite 3 with dorsal hump. Uropod 1 elongate, pe- duncle much longer than rami; rami sub- equal in length (Figs. 32, 33). Uropod 2 shorter than uropod 1, peduncle slightly longer than inner ramus; outer ramus short- er than inner ramus (Figs. 34, 35). Uropod 3, peduncle broad and short; rami subequal in length, elongate, apically obliquely trun- cate (Figs. 36, 37). Telson long and slender, cleft almost to base, with short dorsal spines sparsely distributed in medial row and 1 ter- minal longer spine on each lobe (Figs. 38, 3)9))), Allotype, ovigerous female, body length 9.6 mm. Similar to holotype, but differs from it as follows. Eyes smaller (Fig. 40). Antenna 1, peduncle articles 1—3 longer; flagellum article 1 short (Fig. 41). Antenna 2 much shorter. Mandibles, spine row bear- ing 9 spines. Maxilla 1, inner lobe with more numerous medial setae (Fig. 42). Maxilliped as that of holotype: inner plate VOLUME 111, NUMBER 3 629 ae A. 950 yu p. 50u ¢, 200y Figs. 1-15. Metatiron bonaerensis, new species. Holotype, adult male. 1, Lateral view; 2, 3, Antennae 1, 2; 4, Upper lip; 5, Right mandible; 6, Right molar; 7, Left mandible; 8, Lower lip; 9, Inner plate of maxilla 1; 10, Outer plate of maxilla 1; 11, Palp of maxilla 1; 12, Maxilla 2; 13, Maxilliped; 14, Gnathopod 1; 15, Dactylus of gnathopod 1. Scales: A, Fig. 1; B, Figs. 4-13, 15; C, Figs. 2, 3, 14. 630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 16-30. Metatiron bonaerensis, new species. Holotype, adult male. 16, Gnathopod 2; 17, Dactylus of gnathopod 2; 18, Peraeopod 3; 19, Dactylus of peraeopod 3; 20, Peraeopod 4; 21, Dactylus of peraeopod 4; 22, Peraeopod 5; 23, Dactylus of peraeopod 5; 24, Peraeopod 6; 25, Dactylus of peraeopod 6; 26, Peraeopod 7; 27, Dactylus of peraeopod 7; 28-30, Epimera 1—3. Scales: A, Figs. 16, 18, 20, 22, 24, 26, 28-30; B, Figs. 17, 19, FIN PBI C15 \y lle VOLUME 111, NUMBER 3 with 2 apical tooth-like spines and several submarginal plumose setae; outer plate broad, carrying 3 apical long spines, 6 short, stout medial spines and submarginal setae; palp slender, with 4 articles, article 4 almost as long as article 3, bearing a distal nail (Figs. 43, 44). Peraeopods 3-7 similar, but bearing more setae. Pleonal epimera 1 and 3, posteroventral corners more rounded than male Fully de- veloped oostegites elongate, narrow, mar- ginally setose, attached to coxae 2—5 (Fig. 45). Urosomites 1 and 2 dorsally less elevated than in male; urosomite 3 without dorsal hump (Fig. 46). Telson with numerous lon- ger dorsal spines arranged in row on each lobe; one lobe bearing 2 subapical spines, other lobe with 1 subapical spine (Figs. 47— 49). Paratypic males and females at different stages of maturity; general appearance of pleonites 1—3 and urosomites 1-3 as in ho- lotype and allotype; no morphological vari- ations were observed. Habitat.—The specimens were dredged from sand bottom, at depths of 9.5—-19 m. Fine grain size sand prevailed, but one sam- pling station contained a high percentage of medium grain size sand. No fractions of lime or clay were found (salinity 32.46— 33.05%o0; temperature 14.1—-15.4°C). Etymology.—The species is named in reference to the biogeographical zone, the Bonaerensian district, where the specimens were found. Remarks.—The new species, Metatiron bonaerensis, is assigned to the genus Me- tatiron Rabindranath, 1972, based on the absence of mandibular palp. Barnard (1972) in his review of the family Synoptiidae de- scribed the genus Tiron Liljeborg, 1865, mentioning the presence of a mandibular palp; he made a brief diagnoses of all its known species; some of them had a man- dibular palp while others [like 7. brevidac- tylus (Pillai 1957) and T. tropakis Barnard 1972] the palp was absent, or its condition 631 was unknown. More recently, other authors including Just (1981), Goeke (1982) and Hirayama (1988) have placed new species that lacked a palp on the mandible in Tiron. Barnard & Karaman (1991) diagnosed the genera Metatiron and Tiron, but did not al- locate species to Metatiron; these authors separated both genera by the absence of a mandibular palp and the presence of mid- dorsal tooth on pleonites 1—3 in Metatiron. The new species described in this paper is assigned to Metatiron, primarily because of the absence of a palp on the mandible, and secondarily because of dorsal crenulations on pleonites 1—3. Ledoyer (1979) supported the proposal of Rabindranath (1972) that the lack of the palp in Metatiron was a character of generic value, as did Thomas (1993) in his identification manual for ma- rine amphipods of South Florida. Metatiron bonaerensis is most closely re- lated to Tiron tropakis Barnard, 1972. They resemble each other in their general ap- pearance; they have the accessory eye formed of two separate ommatidia, both lack the palp on the mandible, the peraeo- pods possess stubby dactyli, and the male and female telsons bear similar dorsal spines. Metatiron bonaerensis is easily dis- tinguished from T. tropakis by the laterally smooth pleonites 1—3 which are serrated in the other species, and the forehead protu- berance compared with the evenly rounded forehead in 7. tropakis; in addition, coxa 7 is more quadrate in the new species, and maxilla 1 inner plate is morphologically different in both taxa. Metatiron brevidac- tylus (Pillai 1957) (transfered by Rabin- dranath 1972), M. caecus Ledoyer 1979, M. triocellatus (Goeke 1982) (transfered by Thomas 1993), 7. ovatibasis Hirayama, 1988 and 7. galeatus Hirayama, 1988 also lack a mandibular palp, but they can be sep- arated from the new species as follows: M. brevidactylus has maxilla 1 inner plate small without setae and telson with only one spine at the middle of each lobe; M. caecus, T. ovatibasis and T. galeatus lack accessory eyes, whereas M. triocellatus has 632 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 32 ——= 31 SS asa ee - >= Z q 46 Figs. 31-49. Metatiron bonaerensis, new species. Holotype, adult male. 31 (a—c), Pleonites 1—3; 32, Uropod 1; 33, Ramus tip of uropod 1; 34, Uropod 2; 35, Ramus tip of uropod 2; 36, Uropod 3; 37, Ramus tip of uropod 3; 38, Telson; 39, Apical lobe of telson. Allotype, ovigerous female. 40, Head; 41, Antenna 1; 42, Inner lobe of maxilla 1; 43, Maxilliped; 44, Fourth article of maxilliped; 45, Oostegite; 46, Urosomites 1—3; 47, Telson; 48, 49, Apical lobes of telson. Scales: A, Figs. 31, 32, 34, 36, 38, 41, 45, 47; B, Figs. 33, 35, 37, 39, 42—44, 48, 49: C, Figs. 40, 46. VOLUME 111, NUMBER 3 accessory eyes composed of three omma- tidia, instead of two as in M. bonaerensis. Acknowledgments The material was made available for this study by the Instituto Argentino de Oceano- grafia (IADO) of Bahia Blanca (Buenos Ai- res province), which organized a scientific survey carried out by the R/V El Austral to study the inner continental shelf in the area El] Rincon off the estuary of Bahia Blanca. I am also indebted to Dr. D. Roccatagliata and Lic. M. Torres Jorda from the Univer- sidad de Buenos Aires (UBA), who col- lected the benthic samples and patiently sorted the amphipod material. Literature Cited Barnard, J. L. 1972. A review of the family Synopiidae (=Tironidae), mainly distributed in the deep sea (Crustacea: Amphipoda).—Smithsonian Contri- butions to Zoology 124:1-94. , & G. S. Karaman. 1991. The families and genera of marine gammaridean Amphipoda (ex- cept marine gammaroids).—Records of the Australian Museum, Supplement 13 (part 1):1— 417, (part 2):419-866. Goeke, G. D. 1982. Tiron triocellatus, a new species of amphipod (Gammaridea: Synopiidae) from the western Atlantic and Gulf of Mexico.— Journal of Crustacean Biology 2:148-153. 633 Hirayama, A. 1988. Taxonomic studies on the shallow water gammaridean Amphipoda of west Kyu- shu, Japan. VIII. Pleustidae, Podoceridae, Pris- comilitaridae, Stenothoidae, Synopiidae, and Urothoidae.—-Publications of the Seto Marine Biological Laboratory 33:39-77. Just, J. 1981. Tiron bellairsi sp. n. (Amphipoda, Syn- opiidae) from coral sand in Barbados, with notes on behaviour.—Zoologica Scripta 19: 259-263. Ledoyer, M. 1979. Les gammariens de la pente externe du Grand Récif de Tuléar (Madagascar) (Crus- tacea Amphipoda).—Memorie del Museo Civ- ico de Storia Naturale di Verona (Ila Serie), Se- zione Scienze della Vita 2:1—150. Liljeborg, W. 1865. On the Lysianassa magellanica H. Milne Edwards and on the Crustacea of the sub- order Amphipoda and subfamily Lysianassina found an [sic] the coast of Sweden and Nor- way.—Nova Acta Regiae Societatis Scientia- rum Upsaliensis, Series 3, 6(1):1—38. Pillai, N. K. 1957. Pelagic Crustacea of Travancore. Ii. Amphipoda.—Bulletin of the Central Re- search Institute, University of Travancore 5:29— 68. Rabindranath, P 1972. Three species of gammaridean Amphipoda (Crustacea) from the Trivandrum coast, India—Zoologischer Anzeiger 188:84— OTe Thomas, J. D. 1993. Identification manual for the ma- rine Amphipoda (Gammaridea): I. Common coral reef and rocky bottom amphipods of south Florida. Department of Environmental Protec- tion, Tallahassee, Florida, 83 pp. (unpublished report). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):634—644. 1998. A new genus and species of ‘‘goneplacid-like” brachyuran crab (Crustacea: Decapoda) from the Gulf of California, Mexico, and a proposal for the use of the family Pseudorhombilidae Alcock, 1900 Michel E. Hendrickx Instituto de Ciencias del Mar y Limnologia, Estaci6n Mazatlan, UNAM. PO. Box 811, Mazatlan, Sinaloa 82000, México Abstract.—A new genus and species of crab, Bathyrhombila furcata, are described from localities in the Gulf of California, western coast of Mexico. The new genus is close to Pseudorhombila H. Milne Edwards and belongs to a group of three genera tentatively assigned to the subfamily Pseudorhombi- linae Alcock, 1900 by Guinot in 1969. The affinity of Bathyrhombila, new genus, with other genera of ““Goneplacidae”’ (Euphrosynoplax Guinot, Pseu- dorhombila H. Milne Edwards, Nanoplax Guinot, Oediplax Rathbun, and Cha- cellus Guinot) is discussed, noting that they all represent primitive transitional forms between the cyclometopous and the catometopous abdomen-sternum or- ganization, and a similar “‘xanthoid-goneplacid”’ facies. On the basis of these affinities, it is proposed that these six genera be included into the family Pseu- dorhombilidae Alcock, within the Heterotremata Guinot, 1977. Resiimen.—Se describe un nuevo género y una nueva especie de cangrejo, Bathyrhombila furcata, recolectado en localidades de el golfo de California, costa oeste de México. El nuevo género se parece a Pseudorhombila H. Milne Edwards y pertenece a un grupo de tres géneros tentativamente asignados a la subfamilia Pseudorhombilidae Alcock, 1900 por Guinot en 1969. Se discute la afinidad de Bathyrhombila, nuevo género, con otros género de ““Goneplacidae”’ (Euphrosynoplax Guinot, Pseudorhombila H. Milne-Edwards, Nanoplax Guin- ot, Oediplax Rathbun, y Chacellus Guinot), sefialando que todos ellos repre- sentan formas primitivas de transici6n entre la organizacidn abdomen-esternon cyclometopes y catametopes, y una apariencia similar de tipo “‘xanthoideo- goneplacideo’’. En base a estas afinidades, se propone que estos seis géneros sean incluidos en la familia Pseudorhombilidae Alcock, dentro del grupo de los Heterotremata Guinot, 1977. Two common and widely distributed families of brachyuran crabs, the Xanthidae and Goneplacidae (sensu Balss 1957) have long been recognized as containing hetero- geneous groups of genera (see Guinot 1970, 1977, 1978; Serene 1984; Williams 1984; Martin & Abele 1986). In the last twenty years, the organization of the Xanthidae sensu Balss has been subject to many changes. The concept of a superfamily Xanthoidea proposed by Guinot (1978) em- phasized the sternal position of female openings (‘‘sternitremes’’) coupled with the coxal or coxo-sternal position of male gen- ital openings, thus relating the Xanthoidea with the heterotremateous arrangement (Heterotremata, as defined by Guinot 1977). Guinot (1978) insisted on the fact that among the Xanthoidea, two groups of fam- ilies should be considered: one with male VOLUME 111, NUMBER 3 opening coxal, and another in which the male opening progressively migrates to a coxo-sternal position. In the later group, the evolutionary process is associated with a modification of the facies, which becomes goneplacid-like (Guinot 1978:266). Guinot (1978) also suggested that when all genera of Goneplacidae sensu Balss will have been reviewed, new families might be added to the second, goneplacid-like group of Xan- thoidea (i.e., those genera with a coxo-ster- nal male openings). A group of goneplacid-like brachyuran crabs presently included in the Goneplaci- dae has long been recognized as represent- ing an intermediate step towards the trans- formation of the cyclometopous (heterotre- matous) abdomen-sternum arrangement (male genital opening coxal; abdominal so- mites 1—2 covering entirely the space be- tween the coxa of P5; sternite plate 8 en- tirely covered by these abdominal somites and not visible ventrally) into a catometo- pous arrangement (male opening sternal; sternite plates 7 and 8 widened, ventrally united and visible ventrally; abdominal seg- ments 1—2 reduced, clearly separated from coxa of P5) (Guinot 1969a, 1978, 1979). For Guinot (1969b, 1970) this group of genera represents an evolutionary step to- wards the more advanced catometopous (thoracotrematous) organization in which the sternum occupies an increasingly wider area between the basal abdominal somites and the coxa of P5, and the male opening moves progressively towards a sternal po- sition. Guinot (1970: 1076, 1080) suggested that several of these genera (i.e., Pseudo- rhombila H. Milne Edwards, 1837, Oedi- plax Rathbun, 1893, and possibly Nanoplax Guinot, 1967) could be integrated in the subfamily Pseudorhombilidae Alcock, 1900 pro parte. Guinot (1969b: 721) also described the genus Chacellus Guinot, 1969b, monotypic at that time, which she considered “‘... [a genus with] une organisation trés proche de Vorganisation cyclométopienne et fait sans doute partie des Crabes formant le passage 635 entre Cyclométopes et Catométopes [an or- ganization close to the cyclometopous or- ganization and probably belonging to the crabs linking the cyclometopous to the ca- tametopous].’’ A second species of Chacel- lus was added by Hendrickx (1989a) who, despite of a “rather xanthoid facies,” in- cluded it in the Goneplacidae with a “‘prim- itive catometopous organization.”’ Another genus, Euphrosynoplax Guinot, 1969b was also described by Guinot (1969b:720), to accommodate an undescribed species of crab from Florida: E. clausa Guinot, 1969b. Again, Guinot (1969b) emphasized the primitive catometopous stage of this genus, relatively close to the cyclometopous ar- rangement. A second species of Euphrosy- noplax was recently described by Vazquez- Bader & Gracia (1991) from the Gulf of Mexico. Although these authors did not clearly illustrate sternite eight, they refer to a (what appears as a primitive) catometo- pous organization of their species, E. cam- pechiensis, with a “‘.. . male opening coxal [and] a small portion of sternite 8 not cov- ered by the second abdominal somite.” The present paper deals with a new spe- cies of crab that present morphological sim- ilarities with those in the above cited gen- era. It is herein considered that this new species requires a new genus. Furthermore, the use of the family Pseudorhombilidae Alcock, 1900, is proposed for a group of six genera with a ‘‘xanthoid-goneplacid”’ facies representing primitive transitional forms between the cyclometopous and the catometopous abdomen-sternum organiza- tion. Abbreviations used in this paper are: CW, carapace width; CL, carapace length; P2 to P5, pereiopods; Pll and P12, male first and second pleopods (gonopods), respectively; SEM, Scanning Electron Microscope; EMU, Estacion Mazatlan UNAM, inverte- brate reference collection; SIO, SCRIPPS Institution of Oceanography, invertebrates collection, La Jolla, California, U.S.A.; LACM, Los Angeles County Museum of 636 Natural History, Los Angeles, California, U.S.A. Drawings were made with a camera lu- cida (Fig. 2). Holotype was photographed using a Kodak TMAX 100 ASA black and white film (Fig. 1), and SEM microphoto- graphs of male gonopods were obtained us- ing the classical technique of acetone de- hydrated, gold-palladium coated gonopods extracted from type material (Fig. 3). Bathyrhombila, new genus Diagnosis.—Carapace 1.4 to 1.5 broader than long, anteriorly convex, slightly con- vex and narrower posteriorly; general shape ‘“‘“xanthoid’’. Antero-lateral margin arched, with 4 teeth, excluding the outer orbital tooth which is well-defined and slightly projecting; second and outer orbital teeth fused, forming an almost straight slightly projecting margin; length of this margin al- most half the frontal width. Postero-lateral border converging posteriorly. Regions rel- atively well marked. Front narrow, less than ¥% maximum width of carapace, slightly projecting forwards, with a shallow median depression, margin sinuous, with a well- marked notch between the external corner and the inner orbital tooth. Orbits reduced in size; eyes relatively small. Upper orbital margin slightly concave, with 2 distinct su- tures; lower orbital margin with 2 strong teeth, inner one acute, outer one rounded. Antennal flagellum long, entering orbit; ba- sal article of antenna relatively long, slight- ly oblique, in contact with front; a small apophyse intercalated between basal article and epistome (pterygostomian upper bor- der); palp folding horizontaly. Interanten- nular septum broadly triangular. Third max- illiped with merus about 0.5 times length of ischium, antero-external angle slightly pro- duced; palp articulating at inner distal angle of merus. Chelipeds large and robust, not markedly unequal in large males, subequal in females; carpus with a blunt, moderately large spine at inner angle; pincers very large, fingers flattened, pointed, tips re- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON curved, gap between fingers reduced. A very conspicuous, strong pterygostomian ridge in front of cheliped articulation. Walking legs slender, long, flattened. Ster- num moderately wide, narrowing in front of P1. Abdomen narrow in its base; segment 2 not overlapping coxa of P5. Abdominal segments 3—5 incompletely fused; suture 3— 4 distinguishable, a remaining notch on both extremities; suture 4—5 obsolete, a re- maining notch on both extremities. A small portion of sternite 8 visible, close to artic- ular condyle of P5, not in contact with ster- nite plate 7. A shallow depresion in front of the abdominal telson. Male genital open- ing coxal and gonopod 1 free. Pll long, slender, with slightly curved apex; stiff sub- terminal spines, with stout and long lateral (apical) process and shorter, hook-like api- cal process. PI2 short, sigmoid. Type species.—Bathyrhombila furcata, new species, by original designation and monotypy. Ethymology.—The name of the genus is a combination of rhombila and bathys, to indicate affinities with the genus Pseudo- rhombila H. Milne Edwards and the fact that most specimens were collected in the bathybenthic region. Bathyrhombila furcata, new species Figs 1-3 Material examined.—Holotype, male (CW 24.6 mm; CL 17.5 mm), 05 Jul 1965, station SIO 65-257, La Paz Bay (24°19'N, 110°26’W), Baja California, Mexico, otter trawl, 55-80 m (coll. W. Baldwin) (SIO C- 2116). Paratypes: male (CW 20.3 mm; CL 14.45 mm), 18 Jan 1968, station MV68-I-59, Gulf of California, north of Angel de La Guarda Island (29°41'N, 113°56'W), Baja Califor- nia, Mexico, 566—644 m, otter trawl, R/V T. Washington (coll. C. Hubbs) (SIO C- 5669A). Male, slightly damaged (CW 22.8 mm; CL 16.35 mm), 18 Jan 1968, station MV68-I-59, Gulf of California, north of Angel de La Guarda Island (29°41'N, . VOLUME 111, NUMBER 3 113°56'W), Baja California, Mexico, 566— 644 m, otter trawl, R/V T. Washington (coll. C. Hubbs) (LACM-68-464.1, ex-SIO C-5669). Non-paratypes: 1 soft shell male (cw 25.95 mm; CL 17.95 mm), 1 damaged soft- shell ovigerous female (CW ca. 17.9 mm; CL ca. 13.0 mm), 18 Jan 1968, station MV68-I-59, Gulf of California, north of Angel de La Guarda Island (29°41'N, 113°56'W), Baja California, Mexico, 566— 644 m, otter trawl, R/V TJ. Washington (coll. C. Hubbs) (SIO-5669C). Description.—Carapace wide (CW/CL ratio 1.40—1.46). Front narrow (7.05 mm wide in holotype), sinuous; fronto-orbital width (13.4 mm in holotype) about half the carapace width. Carapace anteriorly convex and mostly covered with small flattened granules, without setae; granules more nu- merous and rounded close to edges and on antero-lateral teeth. Antero-lateral margin with five teeth (including the outerorbital tooth), the posterior three large to medium- sized, conical; second teeth reduced, fused with the outerorbital tooth and forming an almost straight, little projecting margin; outerorbital tooth little produced, distinct. Fifth tooth smaller than the preceeding two; fourth teeth acute, pointing upward; third teeth wider that fourth and fifth, its sides.at a right angle, flattened compared to fourth. Orbital lobe well-marked; upper orbital margin somewhat irregular, with small rounded granules and two conspicuous su- tures (median and lateral); lower orbital margin with granulated inner tooth and out- er lobe, both granulated. Outer orbital tooth well-defined, little projecting. Pterygosto- mian and subhepatic regions granulated. Pterygostomian ridge coarsely granulated. Distal border of merus of third maxilli- ped sinuous, with a marked median concav- ity; antero-external angle little produced; merus coarsely granulated; ischium with more flattened granules, its distal border produced internally in a lobe; palp coarse. Cheliped very strong, long (length of ma- jor cheliped ca. 1.75 CW); claw heavy and 637 long (length of major claw about equal to CW), right claw being slightly higher (right/left claw maximum height ratio 1.09 to 1.12). Merus with granules on anterior and posterior sides, a dorsal row of granules and a blunt superior subterminal angle, pro- duced in a low tubercle. Carpus strong, obliquely subquadrate in dorsal view, sur- face slightly irregular dorsally; clusters of granules arranged in rugae (well defined in the holotype) on outer slope; a blunt, mod- erately large spine at inner angle; a well- defined sulcus parallel to distal border. Ma- nus inflated, smooth (microscopically punc- tated). Fingers long, flattened, pointed, strongly incurving and with recurved tips, gap between fingers reduced; length of dac- tylus of major claw ca. 0.4 times length of claw; dorsal margin of dactylus of major claw almost straight, that of major claw only slightly curved. Cutting edge of dac- tylus of both claws sharp, that of major claw with a strong, projecting subrectan- gular proximal tooth, followed by a series of irregular, smaller teeth; cutting edge of polex with a series of irregular teeth; cut- ting edges of smaller claw with reduced teeth. Pereiopods 2—5 long, slender, flattened; merus covered with dense granules on low- er and upper margin, sides almost smooth; carpus and propodus partly covered with granules on upper margin; a low, longitu- dinal granulated crest on upper margin of carpus; dactylus about same length as prop- odus, with longitudinal rows of setae, tip short, corneous. Pereiopods 2—4 subequal in length (P2 = 1.66 times CW; P3 = 1.69 times CW; P4 = 1.63 times CW), fifth pe- reiopod notably shorter (1.39 times CW). A small portion of sternite eight of male abdomen visible between second and third abdominal somites; first and third somites slightly wider than second, second and third of about the same length and with subacute lateral margins; sixth somite wider than long, sides concave, narrower medially, dis- tal and proximal margin equal; seventh so- mite (telson) as long as sixth, posteriorly 638 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. C-2116). rounded. First somite and lateral portion of second coarsely granulated; other somites with fewer flattened granules or almost smooth. Female gonopores longitudinally oval; opening vertical. First pleopod of male long, slender, bending and slightly curved distally. Two rows of small spines on the shaft; two series of 3 and 5 much longer distal spines on each side of the fold, close to the apex; a strong, spine-like subterminal process, and a terminal, hook-shaped shorter process; a cluster of spines on the side opposite to the fold; an obscure third lobe, covered with tiny spines, in front of the hooked process. Ethymology.—The name of the species refers to the peculiar arrangement of the subapical setae of the male first pleopod, simulating a fork (furca). Remarks.—The smaller male features more marked heterochely, the right claw being about 1.34 times the maximum height Holotype, male, CW 24.6 mm, CL 17.5 mm, dorsal view, La Paz Bay, Baja California, Mexico (SIO of the smallest whereas it is 1.11 in the ho- lotype. The soft-shell male is also the larg- est male available but due to the lack of calcification this specimen has not been used as holotype. The only available female also features a soft-shell, and although basic characteristics match the description of the species, it was not designated as type ma- terial either. The bathymetric range of B. furcata is rather wide; the holotype was taken in trawl between 55 and 80 m, while the rest of the material was obtained in a single trawl from a depth of 566—644 m. Discussion Like several other genera included in the Goneplacidae or “‘Goneplacid-like”’ group (i.e., Pseudorhombila, Nanoplax, Oediplax, and Chacellus), Bathyrhombila represents a primitive evolutionary step towards a ca- tometopous stage, in which an uncovered expanded sternite 8 unites to sternite 7. In VOLUME 111, NUMBER 3 B Fig. 2. 639 Holotype male, CW 24.6 mm, CL 17.5 mm (SIO C-2116). A, dorsal view of carapace; B, dorsal view of abdomen and sternal plates 7—8; C, right (upper) (SIO C-2116) and left (lower) claws, frontal view. the former four genera, the abdominal so- mite 2 is: notably reduced, its lateral margin straight and clearly separated from the coxa of P5 (e.g., Oediplax granulata Rathbun, 1893, type species of the genus; Pseudo- rhombila xanthiformis Garth, 1940; P. oc- todentata Rathbun, 1906); reduced but an- tero-laterally acute, in such a way that the acute corner is close to the coxa of P5 (e.g., Chacellus pacificus Hendrickx, 1989a); or antero-laterally acute and touching the coxa of P5 (e.g., Nanoplax xanthiformis A. Milne Edwards, 1880). In all cases, somite 2 leaves a reduced portion of sternite 8 vis- ible at the basis of coxa of pereiopod 5. In Bathyrhombila the antero-laterally pro- duced corner of somite 2 is almost in con- tact with the coxa of P5. In his study of Bathyplax typhlus oculiferus Miers, 1886, Tavares (1996: 420) note that the size of vissible portion of sternite 8 varies among specimens of a same species; data related to other genera, however, are lacking. Pseu- dorhombila, Nanoplax and Oécediplax are considered by Guinot (1969b, 1970) as po- tential members of a series of “‘Goneplaci- dae”’ related to the Xanthidae, equivalent to the Pseudorhombilinae Alcock, 1900. Among the species of Pseudorhombila, the abdomen-sternum organization itself varies from a primitive step (sternites 7 and 8 ap- pear not in contact in ventral view, male opening coxal: P. xanthiformis) to a more advanced phase [sternites 7—8 in contact on a short distance, in ventral view, displace- ment of the male opening towards a sternal position: P. guadridentata (Latreille, 1828), P. octodentata (Rathbun, 1906), and P. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Microphotographs (SEM) of male first gonopod (P11). A, B and E, E paratype, male, CW 22.8 mm, CL 16.35 mm (LACMNH 68-464.1), slightly damaged, station MV68-I-59, Gulf of California, north of Angel de la Guarda Island (29°41’N—-113°56’W), Baja California, Mexico, detail of extremity; C, D, holotype, male, CW 24.6 mm, CL 17.5 mm (SIO C-2116) distal portion (SEM photographs) (scale: A, EK 100X; B, E, 72; C, D, 25x). ometlanti Vazquez-Bader & Gracia, 1995]; Guinot (1969a, b) is distinguishable in all this would make Pseudorhombila the most the above mentioned genera (in particular advanced genus of all. considering the shape of the cheliped) and The “‘faciés goneplacien” referred to by the pterygostomian ridge, also referred to VOLUME 111, NUMBER 3 by Guinot (1969a, 242) as a “*.. . caractére assez constant [in Goneplacidae], rarement signalé .. .”’ [a rather constant feature, rare- ly reported], is also present in all four gen- era (strong and sharp in Oediplax; strong and granulated in Bathyrhombila; strong to moderate in Pseudorhombila; weak in Cha- cellus). In the case of B. furcata, the general aspect of the crab relates it to species of Pseudorhombila; the carapace is_ similar, notably wider than long, with distinguish- able regions and convex in lateral view. Both genera feature large to very large (Bathyrhombila), heavy chelipeds, although the heterochely is more marked in Pseu- dorhombila. Orbits and eyes are small. Structure of the frontal and antennular regions and of the bucal frame is similar, as is the shape of the third maxilliped, al- though the antero-lateral angle of the merus is not so produced in Bathyrhombila (in this respect, closer to Oediplax). Both genera also feature long (Pseudorhombila) to very long (Bathyrhombila), flattened, slender P2—P5. Bathyrhombila, however, differs from Pseudorhombila in the following characters: the exorbital and second teeth of carapace are fused, forming a wide, almost straight slightly projecting margin (exorbi- tal tooth almost wanting in Pseudorhombi- la); the pterygostomian ridge, in front-of the cheliped articulation, is much stronger in Bathyrhombila; the second abdominal segment is wider in Bathyrhombila; and the structure of the male Pll is strikingly dif- ferent (Fig. 3). Considering the shape of the male gonopods, the slender Pll of Bathy- rhombila is closer to Pll of Chacellus (Pseudorhombila and Oediplax possess a shorter, more massive Pll). Ornamentation of the tip of Pll however, shows affinities with some species of Panopeidae Ortmann, 1893 such as Lophopanopeus frontalis Rathbun (see Martin & Abele 1986: fig. 1N). Although the “‘third’’ lobe (typical of Panopeidae) in Bathyrhombila furcata, new species is hardly distinguishable, the other two processes are strongly developed. In contrast, long subterminal spines (present 641 on the Pll of B. furcata) are also observed on Pll of species of Pseudorhombila and on the type-species of Nanoplax [i.e., N. xanthiformis (A. Milne Edwards, 1880)] (Hendrickx 1995), with a single cluster of subterminal spines in the later. Another genus close to the ‘“‘Pseudo- rhombilid”’ organization is Euphrosynoplax Guinot. The visible portion of sternite 8, however, is smaller in E. clausa (the type- species of the genus) than in Pseudorhom- bila, Bathyrhombila and Oediplax; and is similar in size to sternite 8 of Nanoplax xanthiformis and the two known species of Chacelus. When all these species are compared, they present striking similarities as far as their general shape and aspect is concerned. On the basis of these considerations, and following the suggestion of Guinot (1970; 1080), the use of the family-group name Pseudorhombilidae Alcock, 1900, is pro- posed for those genera of “‘goneplacid-xan- thid”’ crabs. Pseudorhombilidae Alcock, 1900 Pseudorhombilinae Alcock, 1900:286, 292, 297, pro parte. Pseudorhombilinae.—Guinot, 1969b:706; 1971:1080. Type genus.—Pseudorhombila. Included genera.—Bathyrhombila new genus, Chacellus, Euphrosynoplax, Nano- plax, Oediplax, and Pseudorhombila. Definition.—Carapace xanthoid, wider than long, with 3—5 (including outer orbital) antero-lateral teeth. Bucal frame widening anteriorly. Orbits of moderate or reduced size, oval. Chelipeds goneplacids, long, heavy, with long, strongly to moderately in- curving fingers. Pterygostomian ridge (in front of chelipeds) strong to moderate. Ab- dominal somites 3—5 at least partially fused, sutures usually visible. Second abdominal somite reduced, its antero-lateral margin in contact with (anterior angle produced) or separated (margin straight) from basis of coxa of P5. Sternal plate wide and slightly 642 to moderately depressed between P1 (min- imum width between P1 equal to 0.60—0.63 times maximum width between P2). A small to relatively large piece of sternite 8 visible ventrally; sternite 8 not touching sternite 7 in ventral view (Pll coxal) or in contact over a short distance (Pl1 displaced towards a sternal position). Pll long and slender or moderately long and strong; or- namentation variable. P12 short, strongly or moderately sigmoid. Genera.—The family is divided into three groups of genera. Group A includes the most primitive catometopous forms (i.e., Nanoplax, Chacellus, Bathyrhombila and Euphrosynoplax); group B_ includes species with a larger visible piece of ster- nite 8 (Pseudorhombila pro parte and Oed- iplax); group C includes species with a larg- er visible piece of sternite 8 in contact over a short distance with sternite 7 (Pseudo- rhombila pro parte). Remarks.—Among the Pseudorhombili- dae, several species present a subtriangular hiatus between the ischium and the merus of the third maxilliped (e.g., Chacellus pa- cificus; both species of Euphrosynoplax; Oediplax granulata; Pseudorhombila xan- thiformis, P. quadridentata and P. guinotae Hernandez-Aguilera, 1982) while other spe- cies (e.g., Chacellus filiformis Guinot, 1969b; Pseudorhombila ometlanti) possess an anteriorly expanded lobe at the inner an- gle of the ischium that makes contact with the proximal margin of the merus (Hen- drickx 1989a: table 1). The third maxilliped of other species have not been illustrated in the literature and specimens have not been available for examination. In Nanoplax, the partially-fused outerorbital and second teeth are much narrower than in any other genus included in the Pseudorhombilidae, al- though still separated by a shallow notch. The carapace of species of Pseudorhombila and Oediplax features a reduced to uncon- spicuous first anterolateral tooth and the fifth tooth varies from well (e.g., P. octo- dentata) to poorly developed (or obsolete) (e.g., P. quadridentata). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Provisional key to genera of Pseudorhombilidae 1. Distance between outer orbital and first anterolateral teeth much shorter than or- bit width; these tooth partially-fused, separated by a shallow notch. Extremity of male Pll with a strong longitudinally projectine flange... 5.7 eee Nanoplax — Distance between outer orbital and first anterolateral teeth about equal to orbit width 2. Pll of male long, very slender and ta- pering, with only a few minute spines along the shaft). 7.20. 3. eee Chacelus — Pll of male stout, strong, twisted, with median to large spines along the shaft and distal part 3. Outer orbital and first anterolateral teeth coalesced, forming an almost straight slightly projecting margin. Extremity of male Pll with two longitudinal series of vVeby Lone Sspiness sna yee Bathyrhombila — Outer orbital and first anterolateral teeth reduced, separated by a granulated space; granules coarse to minute 4. Merus of MXP3 not produced antero-lat- erally Oediplax — Merus of third maxilliped strongly pro- duced antero-laterally 5. Extremity of male Pll with a strong sub- terminal or lateral upturned flange; a patch of strong spines just below the flange and series of weaker spines along Shaftigesekeiis 0. satus t eee Pseudorhombila — Extremity of male Pll with a lateral flange; no patch of strong subterminal spines below the flange, but a series of moderate size spines along the shaft... PAE Oi emits e. 5.0 Euphrosynoplax Relationships between Pseudorhombili- dae Alcock and the Panopeidae Ortmann are difficult to establish. As emphasized by Guinot (1969a:249, 250, and in lit.) this is due basically to the complexity of the Pan- opeidae sensu lato, of which a first group presents a xanthid facies and coxal male opening (e.g., Eurypanopeus, Panopeus), while a second group presents a goneplacid facies and coxal or coxo-sternal male open- ing (e.g., Cyrtoplax, Glyptoplax). Some VOLUME 111, NUMBER 3 species of the Panopeinae second group (e.g., Glyptoplax pugnax Smith, 1870, the type species of the genus, and G. consagae Hendrickx, 1989b) present a sternum-ab- domen organization more advanced to- wards a catometopous organization, with uncovered section of sternites seven and eight widely in contact and covering a groove through which the penis passes (see Hendrickx 1989b: 653). All members of the Panopeidae, including Glyptoplax, howev- er, feature the typical “‘Panopeid”’ Pll or- namentation. Specimens of the monospecific genera Thalassoplax Guinot and Robertsella Guin- ot were not available during this study. Both genera were briefly described by Guinot (1969b) to accomodate specimens erroneously identified by Rathbun (1918) as Pilumnoplax elata (A. Milne Edwards, 1880). Both genera are very similar primi- tive catometopous and feature distinctive male Pll. Shape of carapace (see Guinot 1969: plate V) of both genera is apparently different from typical pseudorhombilids (carapace more squarish; wider front and orbits). Further studies will be needed to show if these two genera belong to the Pseudorhombilidae or not. Acknowledgments The author thanks W. Newman and L. Spencer, Scripps Institution of Oceanogra- phy, for the loan of the specimens. Com- ments and suggestions provided by D. Guinot, T. Tavares and R. Lemaitre greatly helped to improve the manuscript. A. Es- parza-Haro helped with some of the figures. M. Cordero edited final version of the manuscript. SEM Figure 3 C, D were pre- pared and taken at Scripps Institution of Oceanography and are courtesy of C. Gra- ham; Figure 3A, B, E, F were prepared and taken at the Institut Royal des Sciences Na- turelles de Belgique by E Fiersand J. Cillis. Literature Cited Alcock, A. 1900. Materials for a Carcinological fauna of India. No. 6. The Brachyura Catametopa of 643 Grapsoidea.—Journal of the Asiatic Society of Bengal 69, 2(3):279—456. Balss, H. 1957. Decapoda. Jn Dr. H. G. Bronns Klas- sen und Ordnungen des Tierreichs.—Fiinfter Band, I. Abteilung, 7. Buch, 12. Lief: 1505-— 1672. Garth, J. S. 1940. Some new species of brachyuran crabs from Mexico and Central and South American mainland.—Allan Hancock Pacific Expeditions 5(3):51—127. Guinot, D. 1967. Recherches préliminaires sur les groupements naturels chez les Crustacés Déca- podes Brachyoures. Les anciens genres Micro- panope Stimpson et Medaeus Dana.—Bulletin du Muséum National d’Histoire Naturelle, 2e Sér. 39(2):345-374. . 1969a. Recherches préliminaires sur les grou- pements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae.—Bulletin du Muséum National d’Histoire Naturelle, 2e. Sér. 41(1):241-265. . 1969b. Recherches préliminaires sur les grou- pements naturels chez les Crustacés Décapodes Brachyoures. VII. Les Goneplacidae (suite et fin).—Bulletin du Muséum National d’ Histoire Naturelle, 2e. Sér. 41(3):688—724. . 1970. Recherches préliminaires sur les grou- pements naturels chez les crustacés brachy- oures. VIII. Synthése et bibliographie.—Bulle- tin du Muséum National d’Histoire Naturelle, Paris, 2e. Sér. 5:1063—1090. . 1977. Données nouvelles sur la morphologie, la phylogenése et la taxonomie des Crustacés Décapodes Brachyoures. These de Doctorat d’Etat és Sciences, soutenue le 21 juin 1977 a l’ Université Pierre-et-Marie-Curie. 2 vol. in fol., p. LXV, 1-486, XVI-XXIV, 56 feuilles n-n., 78 Fig., 31 pl., 2 Fig. n.n., 14 tabl. . 1978. Principes d’une classification évolutive des Crustacés Décapodes Brachyoures.—Bul- letin Biologique de la France et de la Belgique 112(3):211—292. . 1979. Données nouvelles sur la morphologie, la phylogenése et la taxonomie des Crustacés Décapodes Brachyoures.—Meémoires du Mu- séum National d’Histoire Naturelle, Paris. Sér. A, Zoologie 112:1—354. Hendrickx, M. E. 1989a. Chacellus pacificus, new spe- cies (Crustacea: Decapoda: Brachyura: Gone- placidae) from the continental shelf of the Gulf of California, Mexico.—Bulletin du Muséum National d’Histoire Naturelle, Paris. Sér. 4, 11, A(1):191—200. 1989b. Glyptoplax consagae new species (Crustacea: Decapoda: Brachyura: Panopeidae) from the Gulf of California, Mexico, with some notes on the distribution of G. pugnax Smith, 1870.—Bulletin du Muséum National 644 d’Histoire Naturelle, Paris. Sér 4, 11, A(3): 649-657. . 1995. Restitution de Pseudorhombila xanthi- formis Garth, 1940, pour Nanoplax garthi Guin- ot, 1969 (Decapoda: Goneplacidae).—Crusta- ceana 68(1):12—20. Hernandez-Aguilera, J. L. 1982. Pseudorhombila guin- otae, un nuevo crustaceo (Decapoda, Gonepla- cidae) en la costa Este de México.—Investiga- ciones Oceanograficas/Biologia 1(4):1—16. Latreille, P. A. 1828. Tourlouroux. Encyclopédie méth- odique: histoire naturelle, entomologie, ou his- toire naturelle des Crustacés, des Arachnides, et des Insectes. Paris 10:345—832. Martin, J. W., & L. G. Abele. 1986. Notes on male pleopod morphology in the brachyuran crab family Panopeidae Ortmann, 1893, sensu Guin- ot (1978) (Decapoda).—Crustaceana 50(2): 182-198. Miers, E. J. 1886. Report on the Brachyura collected by H.M.S. Challenger during the years 1873— 76. In Report Scientific Research Voyage H.M.S. Challenger. Zoology part 49, 17, Lon- don, Edinburgh and Dublin, 1, 362 pp. Milne-Edwards, A. 1873-1881. Etudes sur les Xip- hosures et les Crustacés de la région mexicaine. In Mission scientifique au Mexique et dans 1’ Amérique Centrale. Rech. Zool. Faune Amé- rique Centrale et Mexique, part 5, vol. 1. Paris, Imprimerie Nationale. 368 p. Milne-Edwards, H. 1937. Histoire naturelle des Crus- tacés, comprenant |’anatomie, la physiologie et la classification de ces animaux. Paris II, 532 Pp- Ortmann, A. 1893. Die Dekapoden-Krebse des Strass- burger Museums. VII. Theil. Abtheilung: Brchyura (Brachyura genuina Boas) II. Unter- abtheilung : Cancroidea, 2. Section: Cancrinea, 1. Gruppe : Cyclometopa.—Zoologische Jahr- bticher 7:411—495. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Seréne, R. 1984. Crustacés Décapodes Brachyoures de l’Océan Indien Occidental et de la Mer Rouge, Xanthoidea: Xanthidae et Trapeziidae. Avec un addendum par Crosnier (A.): Carpiliidae et Menippidae.—Faune Tropicale XXIV: 1—400. Rathbun, M. J. 1893. Descriptions of new genera and species of crabs from the west coast of North America and the Sandwich Islands. Jn Scientific results of explorations by the U.S. Fish. Com- mission Steamer “Albatross”. No. XXIV.— Proceedings of the United States National Mu- seum 16(933):223—260. . 1906. Description of a new crab from Dom- inica, West Indies.—Proceedings of the Biolog- ical Society of Washington 19:91. . 1893. The grapsoid crabs of America.—Bul- letin of the United States National Museum 97: 1—461. Smith, S. I. 1870. Notes on American Crustacea. No. 1. Ocypodoidea.—Transactions of the Connect- icut Academy of Arts and Science 2:113—176. Tavares, M. 1996. Sur la validité de Bathyplax typhius oculiferus Miers, 1886 (Decapoda, Brachy- ura).—Crustaceana 69(3):413—423. Vazquez-Bader, A. R., & A. Gracia. 1991. Euphrosy- noplax campechiensis, new species (Crustacea, Decapoda, Brachyura, Goneplacidae) from the continental shelf of southwestern Gulf of Mex- ico.—Bulletin du Muséum National d’ Histoire Naturelle, Paris. 4e. Sér. 13(3—4):433—438. , & . 1995. A new crab species of the genus Pseudorhombila H. Milne-Edwards, 1837 (Crustacea: Decapoda: Goneplacidae).—Pro- ceedings of the Biological Society of Washing- ton 108(2):254—265. Williams, A. B. 1984. Shrimps, lobsters and crabs of the Atlantic coast of the Eastern United States, Maine to Florida. Smithsonian Institution Press, 550 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):645-652. 1998. A new crayfish of the genus Orconectes from the Blood River drainage of western Kentucky and Tennessee (Decapoda: Cambaridae) Christopher A. Taylor and Mark H. Sabaj Center for Biodiversity, Illinois Natural History Survey, 607 E. Peabody Drive, Champaign, Illinois 61820, U.S.A. Abstract.—A new crayfish, Orconectes burri, is described from the Blood River drainage of western Kentucky and Tennessee. The species occurs in small to medium-sized streams with substrates of gravel and sand. Form I males of O. burri differ from all other members of the genus Orconectes in possessing a first pleopod with terminal elements that comprise 22 to 29% of the total length of the pleopod, and a central projection with a tip that terminates at an 90° angle to the main shaft of the pleopod, which is either directly above or slightly overhanging the distal end of the mesial process. Field work conducted in western Tennes- see in the spring of 1996 revealed the pres- ence of an undescribed species of crayfish in the headwaters of the Blood River drain- age. Subsequent field work in western Ken- tucky and Tennessee and examination of material housed at the National Museum of Natural History, Smithsonian Institution, indicated that this undescribed form was endemic to the Blood River drainage. Based on the morphology of the form I male pleo- pod, this new species, described herein as Orconectes burri, is assigned to the Rafi- nesquei group (Fitzpatrick 1987) of the sub- genus Crockerinus Fitzpatrick 1987. Mem- bers of the Rafinesquei group inhabit lotic habitats and are distributed across the west- ern half of Kentucky and extreme southern Illinois. Fitzpatrick (1987) included the fol- lowing as members of the Rafinesquei group: Orconectes bisectus Rhoades, 1944, O. illinoiensis Brown, 1956, O. rafinesquei Rhoades, 1944, O. tricuspis Rhoades, 1944. Orconectes burri, new species Fig. 1, Table 1 Diagnosis.—Body and eyes pigmented. Rostrum slightly concave dorsally, termi- nating in short acumen (see Variation); weak median carina present. Rostral mar- gins thickened; distal halves straight and slightly converging, proximal halves slight- ly convex; terminating in spines (Fig. 1D). Areola 28.8—34.0% (X = 31.0, n = 28, SD = 1.3) of total length of carapace, narrow- est part slightly anterior of midpoint, 4.4— 75 (<= 5-7. n = 28, SD’= 0.9) times as long as wide with 5 to 8 (mode = 6, n = 28, SD = 0.7) punctations across narrowest part (Fig. 1D). One corneous cervical spine on each side of carapace (Fig. 1D). Post- orbital ridges well developed, terminating in corneous spines (Fig. 1D). Suborbital an- gle weakly developed, forming broadly rounded projection. Antennal scale broadest distal to midlength, distal margin at 90° an- gle to lateral margin, thickened lateral mar- gin terminating in large corneous spine (Fig. 1F). Ischia of third pereiopods of males with hooks; hooks overreaching bas- ioischial articulation in form I males only. Chela with 2 rows of tubercles (see Varia- tion) along mesial margin of palm, usually 8 or 9 tubercles in mesial row and 6 or 7 in dorsomesial row; small tufts of setae Over mesial margin of palm, dorsomesial 646 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Orconectes burri, new species: A, Mesial view of first pleopod of form I male; B, caudal view of first pleopods of form I male; C, Lateral view of first pleopod of form II male; D, Dorsal view of carapace; E, Annulus ventralis; EK Dorsal view of right antennal scale; G, Dorsal view of left chela (note- small tufts of setae Over mesial margin of palm, dorsal, dorsomesial, and dorsolateral surfaces not shown). A, D, E G of holotype (INHS 6663); B, topotype (INHS 6662); C, morphotype (INHS 6659); E, allotype (INHS 6664). Scale bars beneath letters equal 1 mm. VOLUME 111, NUMBER 3 Table 1.—Measurements (mm) of Orconectes burri, new species. Holotype Allotype | Morphotype Carapace Total length 26.7 24.1 21.9 Postorbital length 21.3 18.8 16.7 Width 13.9 12.7 10.6 Height 12.2 10.8 9.3 Areola Width 1.5 1.4 1.4 Length 8.7 7.4 6.7 Rostrum Width 3.8 3.5 3.3 Length 6.4 6.0 SES) Chela, left Length, palm mesial margin 8.5 Doll 4.8 Palm width 10.7 7.2 6.0 Length, lateral mar- gin 23.3 15.5 14.2 Dactyl length 12.8 8.3 7.8 Abdomen Width 12.5 12.8 9.2 Length 30.4 27.8 237) and dorsolateral surfaces, and fingers; dor- sal surfaces of fingers with well defined longitudinal ridges (Fig. 1G). First pleopods of form I male symmetrical, extending to posterior edge of bases of second pereio- pods when abdomen flexed. First pleopod of form I male without shoulder on cephalic surface at base of central projection; central projection corneous, constituting 22.2— 29.2% (X = 26.4, n = 9, SD = 2.1) of total length of first pleopod, arched caudodistal- ly, tapering to a pointed tip, tip at 90° angle to main shaft of pleopod and extending to or slightly overhanging distal end of mesial process; mesial process straight and slightly subequal in length to central projection, non-corneous, tapering to an acute tip (Fig. 1A). First pleopod of form II male non-cor- neous, extending to posterior edge of bases of second pereiopods when abdomen flexed; central projection weakly arched caudodistally, mesial process straight and subequal in length; both elements tapering 647 to rounded tips (Fig. 1C). Annulus ventralis immovable, subrhomboidal; cephalic half with wide median trough and 2 caudally di- rected protuberances overhanging centrally located fossa; sinuate sinus running from left corner of fossa to caudal edge (Fig. 1B). Description of holotypic male, form I.— Body slightly depressed dorsoventrally, car- apace wider than abdomen (13.9 and 12.5 mim, respectively). Greatest width of cara- pace larger than height at caudodorsal mar- gin of cervical groove (13.9 and 12.2 mm, respectively). Postorbital carapace length 80.0% of length of carapace. Areola 5.8 times longer (8.7 mm) than wide (1.5 mm) with 6 punctations across narrowest part; length of areola 32.6% of total length of carapace. Rostrum densely covered by se- tiferous punctations, slightly excavated dor- sally, weak carina present; margins thick- ened, distal halves straight and slightly con- verging, terminating in corneous spines, proximal halves slightly convex. Acumen terminating in upturned corneous spine and reaching midpoint of antennular peduncle. Postorbital ridges well developed, terminat- ing in corneous spines. Suborbital angles weakly developed, forming broadly round- ed projections. Cervical spines corneous; dorsal and branchiostegal areas of carapace densely punctate. Abdomen longer than carapace (30.4 and 26.7 mm, respectively). Cephalic section of telson with 1 movable and 1 immovable spine in each caudolateral corner. Protopod- ite of uropod with spine extending over en- dopodite and spine in caudolateral corner extending over exopodite. Caudal margin of cephalic section of exopodite with numer- ous fixed spines (13) and 1 movable spine in caudolateral corner. Lateral margin of en- dopodite terminating in spine; endopodite with prominent median ridge terminating in premarginal spine. Dorsal surfaces of telson and uropods setiferous. Antennal scale broadest distal to mid- length, distal margin at 90° angle to lateral margin, thickened lateral margin terminat- 648 ing in large corneous spine. Right antennal scale 5.5 mm long, 2.3 mm wide. Mesial surface of palm of left chela with 2 rows of tubercles, 9 tubercles in mesial row, 9 tubercles in dorsomesial row, and 3 small widely interspersed tubercles lateral to dorsomesial row. Mesial, dorsal, and lat- eral surfaces of chela covered with numer- ous setiferous punctations; ventral surface with scattered punctations. Dorsal and ven- tral surfaces of finger of propodus with sub- median longitudinal ridges flanked by seti- ferous punctations; basal half of opposable margin with 1 weakly developed tubercle and 4 well developed tubercles; 1 corneous tubercle at midlength between distal-most tubercle and distal tip of finger. Dorsal and ventral surfaces of dactyl with submedian longitudinal ridges flanked by setiferous punctations; basal half of opposable margin with 5 well developed tubercles, first and fourth tubercle from base of dactyl slightly larger than remaining 3. Finger of propodus and dactyl with subterminal corneous tip. Carpus with deep oblique furrow dorsal- ly; mesial margin with 1 large corneous procurved spine just distal to midlength; ventral surface with 1 corneous spine just lateral to midlength of distal margin, 1 Spine just mesial to midlength of distal mar- gin. Dorsodistal surface of merus with 2 corneous spines: ventral surface with 2 large corneous spines just distal to mid- length of ventrolateral margin and mesial row of 7 spines, some corneous; row ter- minating in large corneous spine; small cor- neous tubercle at distolateral corner. Ischi- um with 1 corneous spine just proximal to midlength of mesial margin and 1 large tu- bercle on distal end of mesial margin. Hook on ischium of third pereiopod only; hook simple, overreaching basioischial ar- ticulation and not opposed by tubercle on basis. First pleopod as in Diagnosis, reach- ing to posterior edge of base of second per- eiopods when abdomen flexed. Description of allotypic female.—Differ- ing from holotype as follows. Areola con- stituting 30.1% of length of carapace and PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 5.3 times longer than wide with 7 puncta- tions across narrowest part. Postorbital car- apace length 77.2% of length of carapace. Dorsomesial row of tubercles along palm of left chela with 6 tubercles. Tuft of long se- tae at base of finger of propodus. Ventral surface of left carpus with 2 corneous spines along distal margin. Ventrolateral margin of left merus with mesial row of 5 spines, some corneous. Ventral surface of merus with large corneous spine at disto- lateral corner. Sternum between third and fourth pereio- pods narrowly V-shaped. Postannular scler- ite 1/2 as wide as annulus ventralis (de- scribed in Diagnosis). First pleopod unira- mous, barely reaching caudal margin of an- nulus when abdomen flexed. Description of morphotypic male, form II.—Differing from holotype as follows. Areola constituting 30.6% of length of car- apace and 4.8 times longer than wide. Post- orbital carapace length 77.0% of length of carapace. Dorsomesial row of tubercles along palm of left chela with 8 tubercles. Dorsodistal margin of carpus with 1 cor- neous spine just mesial to carpus/chela ar- ticulation joint. Hook on ischium of third pereiopod not overreaching basioischial articulation. Left fourth and fifth pereiopods detached, right fifth pereiopod detached. First pleopod as described in Diagnosis. Size.—The largest specimen examined is a 33.5 mm total carapace length (CL) form I male. Females (m = 11) range in size from 13.0 to 29.7 mm CL. Form I males (n = 10) range from 14.9 to 33.5 mm CL. Form II males (n = 7) range from 14.0 to 25.2 mm CL. Color.—Dorsal and lateral surfaces of cephalothorax, abdomen, and tail fan dense- ly mottled with tan, light brown, and dark brown patches of varying size. Dorsum with one large laterally elongate dark brown patch just anterior to areola. Ceph- alothorax with dark brown dorsolateral U- shaped saddle connected at caudal margin and extending to midlength of lateral sur- VOLUME 111, NUMBER 3 faces. Saddle discontinuous and mottled an- terior to midlength. Dorsal and lateral sur- faces of chelae, carpus, and merus mottled as cephalothorax and abdomen. Large rounded tubercle at carpus/chela articula- tion joint at dorsoposterior margin of chelae dark blue to black. Fingers of chelae with orange tips followed proximally by wide black bands. Dorsal surfaces of pereiopods tan and mottled with small light brown patches. Ventral surfaces of chelae, cepha- lothorax, and abdomen cream to white. Type locality.—Wildcat Creek at Ken- tucky Hwy. 280, 12.6 km E Murray, Cal- loway County, Kentucky. Holotype was collected from under an exposed tree root mass along the north bank, approximately 5 m downstream of the bridge. The allotype was collected from woody debris that had accumulated just upstream of the bridge. At the time of collection, Wildcat Creek ranged in width from 8—10 m with an av- erage depth of 0.4 m. A small exposed shoal with woody vegetation occurred just upstream of the bridge. Substrate at the type locality was dominated by sand and gravel. Gravel substrates were predominantly found in shallow riffles while sand was found in slower flowing runs and pools. Stream banks were generally steep and well vegetated. Disposition of types.—The holotype, al- lotype, and morphotype are deposited in the Illinois Natural History Survey Crustacean Collection (INHS 6663, INHS 6664, and INHS 6659, respectively), as are the fol- lowing paratypes; 2 ¢ I, 1 ¢ II, and 1 & (INHS 6665). Paratypes consisting of 1 3 I and 2 do II (USNM 260869) and 2 ¢ II, 11 juvenile d, 1 female, and 2 juvenile @ (USNM 148718) are deposited at the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. The localities and dates of collection are provid- ed in the following Range and specimens examined section. Range and specimens examined.—Or- conectes burri, new species, is confined to the Blood River system (Tennessee River 649 Drainage) in western Kentucky and Ten- nessee (Fig. 2). From its headwaters origi- nating in Henry County, Tennessee, the Blood River flows northeasterly across the state line and through Calloway County, Kentucky. This relatively small drainage is best characterized as lowland stream habitat with minimal gradient (Burr and Warren 1986). A large portion of the main channel is embayed at its mouth by Kentucky Lake. Headwaters and tributaries of the Blood River drain Tertiary Porters Creek Clay/ Clayton formations and Cretaceous Mc- Nairy Sand formations while the main channel flows through Quaternary Alluvial deposits. In western Kentucky and Tennes- see the Porters Creek Clay/Clayton and McNairy Sand formations occur as a thin (16-32 km wide) strip of land that roughly coincides with the divide between eastward flowing tributaries of the lower Tennessee River (and the northern flowing Clarks Riv- er) and westward flowing tributaries of the Mississippi River. Extensive collecting in the lower Tennessee River drainage of western Tennessee and Kentucky by the au- thors and a search of holdings in the Na- tional Museum of Natural History have failed to document the presence of O. burri outside the Blood River drainage. In tribu- taries of the Tennessee River south of the Blood River, the species is replaced by Or- conectes (Orconectes) pagei Taylor & Sa- baj, 1997. To the north, O. burri is replaced by a disjunct population of O. (Procericam- barus) durelli Bouchard & Bouchard, 1995. A total of 92 specimens has been exam- ined from the following six locations in Tennessee and Kentucky: TENNESSEE: Henry County: 1) INHS 5815, North Fork Blood River at Blood River Rd., 6.4 km ENE Puryear, 15 Jul 1996 (6 juvenile ¢, 2 2, 10 juvenile 2); 2) INHS 6659, INHS 5822, USNM 260869, Middle Fork Blood River at Mt. Pleasant Rd., 6.4 km E Pur- year, 8 May 1996 (morphotype; 3 6 I, 10 6 Il; 1 6 I, 2 6 II paratypes to USNM); KENTUCKY: Calloway County: 3) INHS 6663, INHS 6664, INHS 6665, INHS 6662, 650 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Known range of Orconectes burri, new species. Type locality denoted by star. 1 = Blood River, 2 = Tennessee River (Kentucky Lake), 3 = Jonathan Creek, 4 = Big Sandy River, 5 = East Fork Clarks River, 6 = North Fork Obion River. Wildcat Creek at KY Hwy. 280, 12.6 kmE km SW Hamlin, 26 Oct 1997 (2 d I, 2 @); Murray (type locality), 26 Oct 1997 (holo- 5) INHS 6669, McCullough Fork at State type; allotype; 2 d I, 1 d Il, 1 2 paratypes; Line Rd., 5.2 km SSW New Concord, 26 431,44 II, 16 2,3 juvenile 2);4)INHS Oct 1997(1 6 1,4 @, 1 juvenile 2); USNM 6667, Beechy Creek at KY Hwy. 121, 7.7. 148718 (2 ¢ II, 11 juvenile ¢, 1 9, 2 ju- VOLUME 111, NUMBER 3 venile @, all paratypes); 6) INHS 5919, Blood River at Grubbs Rd., 0.3 km N KY- TN state line, 31 Mar 1997 (2 6 JI). Etymology.—Named in honor of Dr. Brooks M. Burr, Professor of Zoology and Curator of Fishes at Southern Illinois Uni- versity at Carbondale. Dr. Burr is an avid naturalist with an intense interest in all aquatic organisms. This interest, coupled with his extensive collecting in the creeks and rivers of Kentucky over the past 20 years, has contributed greatly to our knowl- edge of the crayfishes and other aquatic species within the state. Habitat and life-history notes.—Orco- nectes burri, new species, occurs in small to medium-sized streams with substrates of sand and gravel. Within these streams, the species was most commonly encountered in woody debris piles or woody vegetation root masses along stream banks. At site #2 (see Range and specimens examined) the species was collected from under large rip- rap immediately downstream of the bridge. Form I males have been collected in the months of March, May, and October. Ju- veniles were commonly encountered in July. Most O. burri collections were com- posed of two distinct year classes, strongly suggesting a two-year life cycle for the spe- cies. No ovigerous females or females bear- ing young have been collected. Crayfish associates.—The following spe- cies were collected from habitats containing O. burri: Cambarus (Lacunicambarus) di- ogenes Girard, 1852, and Procambarus (Ortmannicus) acutus (Girard 1852). Variation.—Several ontogenetic varia- tions are observed in O. burri, new species, none of which shows any geographic pat- terns of distribution. In smaller individuals the acumen is usually longer, the U-shaped saddle pattern on the cephalothorax is more clearly defined, and fewer tubercles occur along the mesial margin of the palm. In ad- dition to the ontogenetic variation, a few individuals had two or three tubercles lat- eral to the dorsomesial row of tubercles along the mesial margin of the palm, and 651 several females had a sinuate sinus running from the right corner of the fossa of the annulus ventralis. Comparisons.—Orconectes burri differs from all other members of the genus Or- conectes in the shape of the form I male pleopod. The pleopod of O. burri in unique in possessing the following combination of characteristics: terminal elements moderate- ly long, central projection comprising 22 to 29% of total length of pleopod; both ele- ments subparallel to main shaft of pleopod; central projection weakly arched caudodis- tally, tapering to a pointed tip, tip at 90° angle to main shaft of pleopod and either even with or slightly overhanging distal end of mesial process. In addition to the unique shape of the form I pleopod, O. burri, new species, differs from other members of the subgenus Crockerinus by possessing an an- tennal scale that is widest distal to mid- length and with a distal margin forming a 90° angle with lateral margin. Relationships.—The form I male pleo- pod of O. burri is most similar in length and general shape to those of members of the subgenus Crockerinus, and we assign O. burri, new species, to that subgenus. In his subgeneric reorganization of the genus Orconectes, Fitzpatrick (1987) subdivided Crockerinus into five groups: Sanbornii, Marchandi, Propinquus, Rafinesquei, and Shoupi. Following Fitzpatrick’s (1987) hy- pothesis that characters associated with am- plexus are most useful for inferring phylo- genetic relationships, we believe that O. burri, new species, most likely belongs to the Rafinesquei group. Orconectes burri, new species, shares with other members of the Rafinesquei group the following char- acters: central projection of form I male pleopod comprising 22 to 29% of total length of pleopod, distal tip of central pro- jection arched caudally, and distinct trough through cephalic half of annulus ventralis. The placement of O. burri, new species, in the Rafinesquei group also is supported bio- geographically given that it occupies a range closer to those of other members of 652 the Rafinesquei group than to other mem- bers of the subgenus. With the exception of O. rafinesquei, which inhabits the upper Rough River drainage of west-central Ken- tucky, all members of the Rafinesquei group occur in the lower Cumberland, Ten- nessee, and Ohio river drainages of extreme western Kentucky and southern [Illinois (Hobbs 1989). Using morphology alone, we are unable to determine the closest relative of O. burri, new species, because the shape of its form I pleopod is equally different from the pleopods of all other members of the Rafinesquei group. Other characters such as the shape of the rostrum, areola, and chelae offer no additional clues to re- lationships. The rostrum of O. burri, new species, is similar to those of all other mem- bers of the group while the areola and che- lae are unique to the group in that the areola is generally wider and the chelae are stock- ier with shorter fingers. Acknowledgments We are grateful to T. J. Near and L. M. Page for assistance in collecting specimens and to L. M. Page and C. H. Dietrich for reviewing an earlier version of this manu- script. We are also indebted to J. Dexter for photographic assistance and to K. Reed for graciously providing records from and ac- cess to the crustacean collections at the Na- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tional Museum of Natural History, Smith- sonian Institution. Literature Cited Bouchard, R. W., & J. W. Bouchard. 1995. Two new species and subgenera (Cambarus and Orco- nectes) of crayfishes (Decapoda: Cambaridae) from the eastern United States ——Notulae Na- turae 471. Brown, P. L. 1956. A new crayfish of the genus Or- conectes from Illinois.—American Midland Naturalist 56:163—167. Burr, B. M., & M. L. Warren, Jr. 1986. A Distributional Atlas of Kentucky Fishes——Kentucky Nature Preserves Commission Scientific and Technical Series Number 4. Fitzpatrick, J. K, Jr. 1987. The subgenera of the craw- fish genus Orconectes (Decapoda: Cambari- dae).—Proceedings of the Biological Society of Washington 100:44—74. Girard, C. 1852. A revision of the North American Astaci, with observations on their habits and geographical distribution.—Proceedings of the Academy of Natural Sciences of Philadelphia 6: 87-74. Hobbs, H. H., Jr. 1989. An illustrated checklist of the American crayfishes (Decapoda: Astacidae, Cambaridae, and Parastacidae).—Smithsonian Contributions to Zoology 480. Rhoades, R. 1944. The crayfishes of Kentucky, with notes on variation, distribution and descriptions of new species and subspecies.—American Midland Naturalist 31:111—149. Taylor, C. A., & M. H. Sabaj. 1997. A new crayfish of the genus Orconectes from western Tennes- see (Decapoda: Cambaridae).—Proceedings of the Biological Society of Washington 110(2): 263-271. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):653-—662. 1998. Two new species of Erugosquilla from the Indo-West Pacific (Crustacea: Stomatopoda: Squillidae) Shane T. Ahyong and Raymond B. Manning (STA) Department of Marine Invertebrates, Australian Museum, 6 College Street, Sydney South, New South Wales 2000, Australia; (RBM) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. Abstract.—The fifth and sixth species of Erugosquilla to be recognized are E. grahami, new species, from Australia and Taiwan; and E. serenei, new species, from Vietnam. Both species resemble E. woodmasoni (Kemp) in lack- ing distinct tubercles adjacent to the median carina of the telson, and in this feature differ from E. massavensis (Kossmann) and E. hesperia (Manning), both from the western Indian Ocean. The two new species differ from E. woodmasoni in having the anterior margin of the ophthalmic somite trapezoidal rather than rounded in shape; and the outer inferodistal angle of the raptorial merus produced into a blunt angle rather than a spine. Erugosquilla grahami and E. serenei can be distinguished by the length of the prelateral lobe of the telson and the color in life of the antennal peduncle. Erugosquilla Manning, 1995 was erect- ed for species of the Oratosquilla wood- masoni species group. Erugosquilla is dis- tinguished from other species of Oratos- quilla and allies by the combination of the broad carapace, smooth dorsum, suppres- sion of the anterior bifurcation of the me- dian carina of the carapace and the api- cally spinulate anterior margin of the oph- thalmic somite. Manning (1995) recognized Erugos- quilla for four species: E. septemdentata (Ahyong 1994), E. woodmasoni (Kemp 1911), E. massavensis (Kossman 1880), and E. hesperia (Manning 1968). Recent sampling from Australia as part of revi- sionary work on the Australian fauna and restudy of Vietnamese material collected by Raoul Seréne revealed two new species of Erugosquilla, one from Australia and Taiwan, and one from Vietnam, described below. The following abbreviations are used: Al, antennule; A2, antenna; AS, abdom- inal somite; CI, corneal index (CI), 100CL divided by cornea width; CL, car- apace length, measured along the midline, excluding the rostral plate; coll., collector or collected by; ER.V., Fisheries Research Vessel; IM, intermediate; LT, lateral; m, meter(s); MD, median; MG, marginal; mm, millimeter(s); MXP, maxilliped; n, number; NSW, New South Wales; PLP, pleopod; SM, submedian; St, stomatopod catalogue, MNHN, Paris; TL, total length, measured on the midline, from the ante- rior margin of the rostral plate to a line between the apices of the submedian teeth of the telson; TS, thoracic somite; WA, Western Australia. Terminology and size descriptors typically follow the conven- tions of Manning (1969, 1978), supple- mented by some abbreviations proposed by Makarov (1979). All measurements are in millimeters. Type material is deposited in the Austra- lian Museum, Sydney (AM); Muséum Na- tional d’ Histoire Naturelle, Paris (MNHN); National Taiwan Ocean University (NTOU), Keelung; and the National Museum of Nat- ural History, Smithsonian Institution, Washington (USNM). 654 Family Squillidae Latreille, 1803 Erugosquilla Manning, 1995 Erugosquilla grahami, new species Figs. 1, 2, 3A Oratosquilla woodmasoni.—Graham et al., 1993:73 [list; not Oratosquilla woodma- soni (Kemp 1911)]. Material.—Holotype: Australia: AM P4276, 1 3, TL 139 mm, off Patonga, NSW, 32°34’S, 151°17’E, trawled, 7-10 m, coll. S. T. Ahyong. Paratypes: Australia: AM P42767 (to USNM), 1 36, TL 133 mm, type locality, coll. S. T. Ahyong, 29 Jan 1994; AM P42762-66, 3 366, TL 92-141 mm, 2 2? &, TL 102-145 mm, type locality, coll. S. T. Ahyong, 12 Feb 1994; AM P42768-70, 3 36, TL 136-151 mm, type locality, coll. S. T. Ahyong, 12 Feb 1994 Other material.—AM P19332, 2 6¢,TL 115-134 mm, 30 miles south of Carnarvon [24°52’S, 113°38’E], WA, sandy mud with Posidonia and Cymodocea banks, 14—18 m, coll. N. Coleman, 3 Jun 1972; AM P19333, 1 2, TL 123 mm, off Carnarvon, WA, 23 m, coll. A. Nickol, May 1972; AM P41798, 1 3d, TL 134 mm, east of Port Hunter, New- Castles NSW.) 32-555. -157 56 Ee) 6or m: ER.V. Kapala, coll. K. Graham, Jun 1990; AM P41799, 1 ¢, TL 140 mm, southeast of Brunswick Heads, NSW, 28°35’S, 153°34’E, 12-15 m, ER.V. Kapala, coll. K. Graham, 11 Aug 1991; AM P42955, 1 6, TL 125 mm, off Newcastle, NSW, 32°55’S, 151°56’E, 69-73 m, ER.V. Kapala, coll. K. Graham, 3 Dec 1990; AM P42956-58, 1 6, TL 144 mm, 2 22, TL 191-141 mn, off Clarence river, NSW, 30°48’S, 153°02’E, 22-30 m, ER.V. Kapala, coll. K. Graham, 5 Nov 1991; AM P42949, 1 2°, TL 94 mm, Port Jackson, NSW, 33°55’S, 151°15’E, coll. M. Beatson, 7 Mar 1994; AM P42950, 1 2, TL 74 mm, Port Jackson, NSW, 33°55’S, 151°15’E, coll. M. Beatson, 7 Mar 1994; AM P42951-54, 2 66, TL 64-116 mm, 2 2 2, TL 122-146 mm, Port Jackson, NSW, 33°51’S, 151°15’E, coll. M. Beatson, 7 Mar 1994. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Taiwan: NTOU, 1 36, TL 91 mm, western Taiwan, 50 m, on sand, commercial trawler, 6 Jul 1996. Description.—Size large, TL of adults to at least 150 mm. Dorsal surface smooth, polished. Eye large, not extending beyond A1 pe- duncle segment 1; cornea strongly bilobed, set obliquely on stalk; CI 295-403 (holo- type 373). Ophthalmic somite anterior margin trap- ezoid, with median spinule. Ocular scales separate, subtruncate. Al peduncle subequal to CL. Al somite dorsal processes trianguloid, apices acute, directed anterolaterally. A2 scale slender, length 0.5—0.7 CL. Rostral plate broader than long, trape- zoid; lateral margins upturned, convergent, straight; apex truncate; dorsal surface lack- ing MD carina. Carapace anterior width 0.6 CL; antero- lateral spines not extending beyond base of rostral plate; MD carina very low, indis- tinct; anterior bifurcation absent; with nor- mal complement of carinae (MD, SM, LT, MG); posterior median projection very low or absent. Raptorial claw dactylus with 6 teeth, out- er margin sinuous, lacking basal notch or lobe; carpus dorsal carina irregular] tuber- culate; propodus opposable margin evenly pectinate, distal margin lacking stout tooth; merus outer inferodistal angle at most a blunt angle. Mandibular palp 3-segmented. MXP1-—4 each with epipod. MXP5 basal segment with ventrally directed spine. TS6—8 each with SM and IM carinae, former indistinct. TS5 lateral process bi- lobed, anterior lobe a slender spine directed anterolaterally, posterior lobe short, trian- gular, apex acute, directed laterally. TS6 lateral process bilobed; anterior lobe much smaller than posterior, slender, apex blunt; posterior lobe broad, triangular, an- terior margin straight, apex acute. TS7 lateral process bilobed, anterior lobe much smaller than posterior, latter lobe VOLUME 111, NUMBER 3 655 Fig. 1. Erugosquilla grahami, new species, male holotype, TL 139 mm, AM P42761. A, Anterior part of body; B, Anterior margin of ophthalmic somite, enlarged; C, Lateral processes of TS5—7; D, AS6, telson, and right uropod. (Setae omitted). Scale: A,C,D, 5 mm; B, 2.5 mm. 656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs 2: of MXP1—5. A, MXP1; B, MXP5: C, MXP4; D, MXP3; E, MXP2. (Setae omitted). Scale: A—D, 2.5 mm; E, 5 mm. broad, triangular, anterior margin straight, apex blunt. TS8 anterolateral margin quadrate, apex blunt; sternal keel rounded. AS1-—5 each with normal complement of carinae (SM, IM, LT, MG). SM carinae low, indistinct; divergent on ASS. AS6 with dis- tinct SM, IM, and LT carinae; with ventro- lateral spine anterior to uropodal articula- tion. Abdominal carinae spined on the fol- lowing somites: SM 5—6; IM 3-6; LT (2)3— 6; MG 1-5. Male PLP1 endopod with hook process approximately half length of tube process; apex blunt. Telson flattened, subquadrate, slightly Erugosquilla grahami, new species, male holotype, TL 139 mm, AM P42761. Distal three segments broader than long; with 3 pairs of primary marginal teeth, apices fixed; prelateral lobe subequal to or longer than margin of LT tooth; denticles SM 3—4, IM 8-10, LT 1, rounded, each with dorsal tubercle; carinae of MG teeth slightly inflated in adult males; MD carina interrupted proximally, armed posteriorly with apical spine overhanging 2 blunt tubercles; posteriorly convergent, pit- ted groove present on each side of MD, lat- eral margin raised, irregular, but not formed by distinct tubercles; dorsolateral surface with curved rows of shallow pits; ventral surface with postanal carina. Uropod protopod terminating in 2 slen- der spines, dorsally and ventrally carinate, VOLUME 111, NUMBER 3 Fig. 3. 657 Endopod of first male PLP (gonopod). A, Erugosquilla grahami, new species, male holotype, TL 139 mm, AM P42761; B, E. serenei, new species, male paratype, TL 125 mm, AM P12151. (Setae omitted.) Scale: 2.5 mm. inner longer, unarmed dorsally excepting dorsal spine above proximal exopod artic- ulation; with minute ventral spine anterior to endopod articulation; protopod inner margin crenulate, terminal spines with small, rounded lobe on outer margin of in- ner spine, margin concave. Uropod exopod proximal segment un- armed dorsally; outer margin with 7—9 (usually 8) sharp, graded, movable spines, distalmost not exceeding midlength of dis- tal segment; distal margin with slender ven- tral spine; exopod distal segment longer than proximal segment; endopod unarmed dorsally, with 1 dorsal and 1 ventral carina. Color in life.—Base color white; overall dorsal surface pale gray-green. Margin of rostral plate, dorsal carinae and gastric grooves of carapace, posterior margin of carapace and thoracic somites, SM carinae of thoracic and abdominal somites, red. Al peduncle with alternating blue and yellow- orange banding. A2 scale blue proximally, yellow distally. Meral-carpal articulation of raptorial merus yellow. Meral depression pale yellow. IM and LT carinae of AS5—6 dark green; apices of spines red. AS2, 5 with dark rectangular median patch. Dorsal surface of telson maroon; carinae and den- ticulate dorsal tubercles dark green; denti- cles and marginal teeth white. Proximal segment of uropodal exopod dark blue; dis- tal segment blue proximally, yellow distal- ly. Endopod blue; distal tip yellow. No col- or differences were noted between the sex- es. Etymology.—tThis species is named for Mr. Ken Graham, NSW Fisheries, who col- lected much of the material used in this study. Measurements.—Holotype: AM P42761, male, TL 139 mm, CL 26.5 mm, cornea width 7.1 mm. Paratypes: Males (n = 7), TL 92-151 mm; females (n = 2), TL 102 and 145 mm. Other material: males (n = 9) 64-151 mm; females (n = 7) TL 74-146 mm. Remarks.—Erugosquilla grahami, new species, is morphologically most similar to E. woodmasoni (Kemp 1911) in the rela- 658 tively short rostral plate and length of the prelateral lobes of the telson (subequal to or longer than margin of LT). The two spe- cies differ in the shape of the anterior mar- gin of the ophthalmic somite (trapezoid in E. grahami and broadly rounded in E. woodmasoni), abdominal carination (E. grahami generally has fewer posteriorly armed lateral carinae), and the condition of the outer inferodistal angle of the raptorial merus (armed in E. woodmasoni). The two species have similar general coloration, in- cluding blue uropods, but may be distin- guished by coloration of the Al peduncles (red-maroon in E. woodmasoni, banded blue and yellow-orange in E. grahami). Differences between E. grahami and E. ser- enei are noted below, under the account of the latter species. Both E. woodmasoni and E. grahami are known from New South Wales, but the lat- ter is more common. Like most squillids, E. grahami inhabits shallow coastal waters and constructs burrows in soft level sub- strates. The present specimens were col- lected in depths of 7—73 m (but usually less than 30 m), over sandy mud and vegetated sand (Cymodocea and Posidonia). The known distribution of E. grahami (i.e., Taiwan and Australia) is discontinu- ous, but likely reflects limited sampling ef- fort in intermediate localities. The disjunct Australian distribution of FE. grahami sug- gests that it probably occurs throughout northern Australian waters. The Taiwanese specimen agrees in almost all respects, in- cluding live coloration, with Australian ma- terial, except that the TS7 lateral process anterior lobe is sharper than in most Aus- tralian specimens. Distribution.—Taiwan and Australia, from northern New South Wales, south to Port Jackson, and Carnarvon, Western Aus- tralia. Erugosquilla serenei, new species Figs. 3B, 4 Squilla massavensis.—Seréne, 1951:fig. 2 [Vietnam]; 1953:507 [Vietnam]; 1954:6, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 8, 54, 60-62, 87, pl. 3, figs. 5—8 [Nha- trang (12°15'’N, 109°12’E) and Cauda (12°11-13'N, 109°13-16’E) bays, Viet- nam]. [Not Squilla massavensis Koss- mann 1880.] Erugosquilla hesperia.—Manning, 1995: 24, 198, figs. 121, 122, 1234) scam 35 [color] [Nhatrang and Cauda bays, Vi- etnam]. [Not Squilla hesperia Manning 1968.] Material.—Holotype: MNHN-St 1940, 1 3, TL 125 mm, Vietnam, Oceanographic Institute of Nhatrang, opposite the labora- tory, coll. R. Seréne, Nov 1948. Paratypes: USNM 266696, 1 3, TL 126 mm, Vietnam, coll. R. Seréne, 20 May 1949; same data, USNM 266693, 1 2, TL 124 mm.—USNM 266695, Nhatrang Bay, 1 2, TL 103 mm, coll. R. Seréne, 21 May 1949.—USNM 277646, 1 2, TL 142 mm, Oceanographic Institute of Nhatrang; AM P12151, 1 3d, 126 mm, same locality, depth 25 m, coll. R. Seréne, 23 Nov 1949; AM P51049, 1 2, 116 mm, same locality, coll. R. Seréne, 23 Nov 1949. Description.—Size large, TL of adults to at least 140 mm. Dorsal surface smooth, polished. Eye large, extending to or slightly over- reaching Al peduncle segment 1; cornea strongly bilobed, set obliquely on stalk; Cl 325-380 (holotype 349). Ophthalmic somite anterior margin trap- ezoidal, with small median spinule (broken in Fig. 4A). Ocular scales separate, sub- truncate. A1 peduncle slightly shorter than CL. Al somite dorsal processes triangular, apices acute, directed anterolaterally. A2 scale slender, length 0.7 CL. Rostral plate appearing elongate, length and width at base subequal; lateral margins upturned, convergent anteriorly, straight; apex truncate; dorsal surface lacking MD carina. Carapace anterior width 0.6 CL; antero- lateral spines not extending beyond base of rostral plate; MD carina distinct posterior to VOLUME 111, NUMBER 3 659 Fig. 4. Erugosquilla serenei, new species. A-D, female paratype, TL 103 mm, USNM 266695; E, female paratype, TL 124 mm, USNM 266693. A, Anterior part of body; B, Carpus of raptorial claw; C, Lateral processes of TS5—8; D, AS5—6, telson, and right uropod; E, anterior margin of ophthalmic somite. (Setae omitted). Scale: A-D, 5 mm; E, 2.5 mm. 660 dorsal pit, anterior bifurcation absent; with normal complement of carinae; posterior median projection very low. Raptorial claw dactylus with 6 teeth, out- er margin sinuous, slightly inflated, lacking basal notch or lobe; carpus dorsal carina ir- regularly tuberculate; propodus opposable margin evenly and finely pectinate, distal margin lacking sharp tooth; merus outer in- ferodistal angle a low, obtuse projection. Mandibular palp 3-segmented. MXP1—4 each with epipod. MXP5 basal segment with low, ventrally directed spine. TS6—8 each with SM and IM carinae, SM carinae indistinct. TS5 lateral process bilobed, anterior lobe a slender spine di- rected anterolaterally, posterior lobe short, triangular, apex acute, directed laterally. TS6 lateral process bilobed, anterior lobe much shorter than posterior, slender, ante- rior margin convex, apex acute; posterior lobe broad, triangular, anterior margin straight, apex acute. TS7 lateral process bilobed, anterior lobe much smaller than posterior; latter broad, triangular, anterior margin straight, apex blunt. TS8 anterolateral margin quadrate, apex blunt; sternal keel rounded. AS1-5 each with normal complement of carinae (SM, IM, LT, MG); SM carinae dis- tinct, subparallel or slightly divergent pos- teriorly. AS6 with distinct SM, IM, and LT carinae; with minute ventrolateral spine an- terior to uropod articulation. Abdominal ca- rinae spined on following somites: SM (4) 5—6, IM (2) 3-6, LT 1-6, MG 1-5. Male PLP1 endopod with hook process approximately half length of tube process; apex blunt. Telson flattened, subquadrate, length and width subequal, with 3 pairs of primary marginal teeth, apices fixed; prelateral lobe shorter than margin of LT tooth; denticles SM2-3, IM7-9, LT1, rounded, each with dorsal tubercle; carinae of MG teeth at most slightly inflated in adult males; median ca- rina interrupted proximally, armed posteri- orly with apical spine overhanging 2 blunt PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tubercles; with single, submedian, curved, irregular carina on each side, converging posteriorly under base of median spine, and dorsolateral surface with curved rows of shallow pits; ventral surface with postanal carina. Uropod protopod terminating in 2 slen- der spines, dorsally and ventrally carinate, inner longer; unarmed dorsally excepting dorsal spine above proximal exopod artic- ulation; with minute ventrolateral spine an- terior to endopod articulation; protopod in- ner margin crenulate, terminal spines with very small, rounded lobe on outer margin of inner spine, margin concave. Uropodal exopod proximal segment un- armed dorsally; outer margin with 8 sharp, graded, movable spines, distalmost not ex- tending to midlength of distal segment; dis- tal margin with slender ventral spine; exo- pod distal segment subequal to proximal segment; endopod unarmed dorsally, with 1 dorsal and 1 ventral carina. Color in life-—Basic color tan. Margin of rostral plate, dorsal carinae and gastric grooves of carapace, carinae of TS and AS, median carina of telson, and some carinae of uropod, red. Al peduncle tan, not mark- edly banded; A2 scale tan. Raptorial dac- tylus ivory white. AS2,5 each with dusky, rectangular dorsal patch. Uropod exopod proximal segment light blue, with some white anteriorly, grading to dark blue dis- tally, proximal segment bluish mesially, white laterally. Uropod endopod reddish mesially, tan laterally. Etymology.—Named for the late Raoul Serene, who first recorded it from Vietnam. Measurements.—Holotype: MNHN-St 1940, male, TL 125 mm, CL 26.2 mm, cor- nea width 7.5 mm. Paratypes: Males (n = 2), both TL 126 mm; females (n = 4) TL 103-142 mm. Remarks.—Erugosquilla serenei resem- bles E. grahami and differs from E. wood- masoni (Kemp) in having the anterior mar- gin of the ophthalmic somite trapezoidal in shape rather than evenly rounded and a blunt angle or lobe rather than a sharp spine VOLUME 111, NUMBER 3 at the outer inferodistal angle of the merus of the raptorial claw. Erugosquilla serenei differs from E. gracilis in having a narrow- er rostral plate; the prelateral lobe of the telson shorter than the LT margin; and, the antennal peduncle uniformly colored in life rather than banded with blue and yellow- orange. Erugosquilla woodmasoni and its four congeners that have six teeth on the dac- tylus of the claw all also have a groove on each side of the MD carina of the telson that converges under the posterior spine of the MD carina. In E. woodmasoni, this groove is shallow and lacks a flanking ca- rina or row of distinct tubercles; the groove is much more distinct in the other four spe- cies. Both E. hesperia and E. massavensis have distinct tubercles flanking this groove, one row in the former, two rows in the lat- ter. In both of the new species the row of tubercles is replaced by an irregularly tu- berculate carina. Acknowledgments We thank Nguyen Ngoc-Ho, Muséum National d’Histoire Naturelle, Paris, for providing information on the holotype of E. serenei, and Tin-Yam Chan, National Tai- wan Ocean University, for allowing us to work with the specimen of E. grahami from his collections. Figure 4 was prepared by Lilly King Manning. This study was par- tially supported by an Australian Postgrad- uate Award to S.T:A. from the Australian Research Council and administered by the University of New South Wales. The sup- port of the Smithsonian Marine Station at Fort Pierce for Manning’s studies of sto- matopods is gratefully acknowledged. This is contribution #446 from that program. Key to Species of Erugosquilla 1. Dactylus of claw with 7 teeth; rostral plate as long as broad, apex evenly OUNCE AEN ous « Seeks Siactah. aa E. septem- dentata (Ahyong 1994) [Indonesia] — Dactylus of claw with 6 teeth; rostral plate usually apically flattened 661 2. Telson median carina flanked by groove on each side, convergent posteriorly, with or without an irregular lateral cari- na, lacking distinct tubercles ......... 3 — Telson median carina flanked by one or more rows of tubercles 3. Ophthalmic somite anterior margin broadly rounded, usually with apical spi- nule; outer inferodistal angle of raptorial merus produced to a blunt spine; A2 pe- duncle red-maroon in life............ E. woodmasoni (Kemp 1911) [western Pacific to western Indian Ocean] — Ophthalmic somite anterior margin trap- ezoid, with apical spinule; outer infer- odistal angle of raptorial merus produced to at most a blunt angle; A2 peduncle not red-maroon in life 4. Telson prelateral lobe subequal to or lon- ger than margin of lateral tooth; A2 pe- duncle banded with blue and yellow-or- ange pin like wee eae E. grahami, new species [Australia and Taiwan] — Telson prelateral lobe shorter than mar- gin of lateral tooth; antennal peduncle uniformly colored E. serenei, new species [Vietnam] 5. Rostral plate lateral margins sinuous; tel- son with 2 rows of tubercles flanking median carina E. massavensis (Kossmann, 1880) [Red Sea and western Mediterranean] — Rostral plate lateral margins straight; tel- son with 1 row of tubercles flanking me- dianicanilaunnr secre E. hesperia (Manning 1968) [western Indian Ocean] Literature Cited Ahyong, S. 1994. Oratosquilla septemdentata n. sp. (Crustacea: Stomatopoda: Squillidae), a new species of deep water stomatopod from Hal- mahera, Indonesia.—Records of the Australian Museum 46:1—4. Graham, K. J., G. W. Liggins, J. Wildforster, & S. J. Kennelly. 1993. Relative abundances and size compositions of prawns and by-catch species on New South Wales prawn grounds during sur- veys V—VIII (May 1991—May 1992).—Kapala cruise report 112:1—73. Fisheries Research In- stitute, NSW. Kemp, S. 1911. Preliminary descriptions of new spe- cies and varieties of Crustacea Stomatopoda in 662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the Indian Museum.—Records of the Indian with accounts of two new genera and nine new Museum 6(2):93—100. species (Crustacea: Stomatopoda: Squillidae).— Kossmann, R. 1880. Malacostraca.—Zoologische Er- Smithsonian Contributions to Zoology 272:1— gebnisse einer im Auftrage der k6niglichen 44. Akademie der Wisssenschaften zu Berlin aus- . 1995. Stomatopod Crustacea of Vietnam: The gefiirten Reise in aie Kiistengebiete des Rothen legacy of Raoul Seréne.—Crustacean Research, Meeres 2(1):1—140, pls. 1-15. Special Number 4:1—339. The Carcinological Makarov, R. R. 1979. A collection of stomatopod crus- Society of Japan, Tokyo. taceans of the genus Clorida Eydoux & Souley- Serene, R. 1951. Sur la circulation d’eau a la surface et, 1842, from Tonkin Bay, Vietnam.—Crusta- du corps des Stomatopodes.—Bulletin de la So- ceana 37(1):39-56. . ciété zoologique de France 76(3):137—143, pl. 1. Manning, R. B. 1968. Stomatopod Crustacea from . 1953. Sur la collection des Stomatopodes de Madagascar.—Proceedings of the United States l'Institut Océanographique de 1 Indochine.— National Museum 124:1-61. Proceedings of the Seventh Pacific Science . 1969. Stomatopod Crustacea of the western Congress, 1949 4(Zoology):506—508. Atlantic.—Studies in Tropical Oceanography, . 1954. Observations biologiques sur les Sto- Miami 8:1—380. matopodes.—Meémoires de 1’Institut Océano- . 1978. Further observations on Oratosquilla, graphique de Nhatrang 8:1—93, pls. 1—10. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):663—673. 1998. Two new species of Macrobiotidae (Tardigrada: Eutardigrada) from the United States of America, and some taxonomic considerations of the genus Murrayon Roberto Guidetti Department of Animal Biology, University of Modena, Italy Abstract.—Two new species of eutardigrades, Macrobiotus nelsonae and Murrayon stellatus, collected in Tennessee, U.S.A., are described. Macrobiotus hibernicus and Macrobiotus dianeae, related to M. stellatus, are re-examined and ascribed to the genus Murrayon on the basis of the claw type and the presence of a ventral hook on the strengthening bar. A dichotomous key to the genus Murrayon is proposed. Thulin (1928), Marcus (1936) and Ra- mazzotti (1962), in their systematic mono- graphs on tardigrades, included all eutardi- grades (except Milnesium) in the family Macrobiotidae and all those with so-called ‘symmetric’ claws, i.e., with respect the median plane of the leg, in the genus Ma- crobiotus. Pilato (1969a, 1969b) proposed to separate the eutardigrades in different families on the basis of the claw structure, ascribing to Macrobiotidae only the genera characterized by ‘symmetric’ claws, i.e., Macrobiotus and Pseudodiphascon (the lat- ter erected as a subgenus by Ramazzotti in 1965). The status of this family remained almost unchanged until the 1980s, when new taxonomic characters, related to the bucco-pharyngeal apparatus and to the claws, were considered for the identifica- tion of the genera (Schuster et al. 1980, Bertolani 1981, Pilato 1981, Bertolani & Kristensen 1987). Today 11 genera are placed in the Macrobiotidae. Moreover, the number of known species has been in- creased remarkably due to a more in-depth analysis and the use of a greater number of characters. Within the genus Macrobiotus, for example, is the so-called ‘hufelandi’ group studied by light microscopy and Scanning electron microscopy (SEM) by Biserov (1990a, 1990b) and by Bertolani & Rebecchi (1993), who identified several new species within this group with so- called high intra-specific variability. Within the genus Macrobiotus, natural groups of species sharing similar morphology of the animals and/or of spermatozoa and eggs were identified, which led to the hypothesis of further possibilities of subdivision at the genus or subgenus level (Guidi & Rebecchi 1996). Tardigrades from new material collected in Tennessee (U.S.A.) and comparisons with known related species have expanded our knowledge of the systematics of Ma- crobiotidae, one of the larger and more complex families of eutardigrades, leading to the erection of two new species and the redesignation of two other species. Methods Specimens from 44 samples of beech leaf litter (Fagus grandifolia Ehrb.) collected in fall 1995, winter 1995-1996, and spring 1996 on Roan Mountain (Southern Appa- lachians, Carter County, Tennessee, U.S.A.) at different altitudes (1200 m, 1500 m and 1650 m asl) were examined. Tardigrades and their eggs were preserved for SEM ex- aminations or directly mounted in polyvi- nyl-lactophenol or in Hoyer’s medium, or stained with acetic carmine and subsequent- 664 ly mounted in Faure-Berlese’s mounting medium for the light microscopy (LM) ob- servations. Light microscopy observations were done with phase contrast and differ- ential interference contrast (DIC) under oil immersion (100). The specimens prepared for SEM were fixed in absolute ethanol, critical point dried in liquid CO, and sput- ter-coated with gold-palladium. All the specimens were examined using a Philips SEM 500 at the ‘Centro Interdipartimentale Grandi Strumenti’ of the University of Mo- dena. As comparison material, several speci- mens were used: Murrayon pullari (Murray 1907) from Monte Rondinaio, Emilia Ro- magna, Italy (from Bertolani’s collection); Murrayon hastatus (Murray 1907) from Angerfjrden, Sweden (from Kristensen’s collection); Macrobiotus hibernicus Murray 1911 from Valtellina, Lombardia, Italy (from Pilato’s collection), from Valico of Gran San Bernardo, Valle d’ Aosta, Italy, from Val Piantonetto, Piemonte, Italy (both from Maucci’s collection; the latter pre- sented to him by C. Robotti) and from God- havn, Greenland (from Kristensen’s collec- tion); Macrobiotus dianeae Kristensen 1982 from Unartog, Greenland (paratypes from Kristensen’s collection), from Monte Rondinaio, Emilia Romagna, Italy (from Bertolani’s collection); a paratype and an egg of Macrobiotus aviglianae Robotti 1970, and the holotype and an egg of Ma- crobiotus pallarii Maucci 1954 (both from Maucci’s collection). Macrobiotus nelsonae, new species (Figs. 1-5, Table 1) Etymology.—The species is dedicated to Dr. Diane R. Nelson, professor at East Ten- nessee State University, whose hospitality and help allowed this work. Paratypes.—566 paratypes and 322 eggs mounted in polyvinyl-lactophenol, Hoyer’s medium and Faure-Berlese (permanent slides), 3 paratypes and 3 eggs on stubs for SEM observation. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Holotype.—Adult male 786.1 wm in length, collected in January 1996, mounted in polyvinyl-lactophenol, slide labeled as SNO5a-S10 (Fig. 1A). Type locality.—Roan Mountain (Carter County, Tennessee) on the north-facing slope at elevations between 1200 and 1650 m asl, in leaf litter from beech trees. Diagnosis.—Animals of large size. Smooth cuticle with pores. Eye-spots ante- rior. Wide buccal tube; evident buccal ar- mature, with two consistent bands of teeth and with transverse crests. Two rod-shaped macroplacoids and microplacoid. Claws of the ‘hufelandi’ type, with lunulae. Eggs laid freely, with big conical reticulated process- es inserted onto an areolate surface. Description.—Length from 196.0 up to 925.4 wm. Colorless, although the largest animals may have a pink coloration. Eye- spots in anterior position. Smooth cuticle, with oval cuticular pores not uniform in size and irregularly distributed, much more visible in the anterior and posterior part of the animal. In some animals a uniform and fine punctuation on the dorsal cuticle is vis- ible, much more identifiable in recently pre- pared specimens (Fig. 2). Ten evident per- ibuccal lamellae. Buccal armature charac- terized by an anterior band of small teeth (mucrones), followed by a large posterior band of larger teeth, whose dimensions in- crease caudally (Figs. 1C, D). Three trans- verse crests, the dorsal ones longer than the ventral; in some animals the median ventral crest is subdivided in two or seldom into more segments of variable length. Wide buccal tube. Rounded pharyngeal bulb (ra- tio length/width 1.1—1.3:1) with large apophyses, two rod-shaped macroplacoids and an evident microplacoid. First macro- placoid the longest, with an evident nar- rowed midportion, the second with a sub- terminal constriction (Fig. 1B). Massive claws, of ‘hufelandi’ type (Y-shaped; Figs. 1E, F), the claws of the front legs the small- est and those of the hind legs the largest. Main branch of the outer claw slightly lon- ger than that of the inner one, both bearing VOLUME 111, NUMBER 3 665 Fig. 1. Holotype of Macrobiotus nelsonae sp. n. A, Habitus (ventral view); B, Bucco-pharyngeal apparatus (frontal view); C, Buccal armature (ventral view); D, Buccal armature (dorsal view); E, Claws of the front legs; K Claws of the hind legs; G, Egg. H, Particular of the egg. Scale bars, 100 wm for A; 20 wm for B, G; 10 pm for C, D, E, FE H. 666 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 2-5. Macrobiotus nelsonae sp. n. 2, Dorsal cuticle with pores (arrow) and fine punctuation (DIC); 3, Egg in toto (SEM); 4, Particular of the egg (Phase contrast); 5, Particular of the egg (SEM). Scale bars, 10 1m for Figs. 2, 4, 5; 20 pm for Fig. 3. evident accessory points. Small lunulae at the base of the claws of the first three pairs of legs. Lunulae surrounded by fine teeth in the largest animals. Lunulae of the hind claws large, even larger on the posterior claws, and with teeth of irregular dimen- sions in the largest animals. Three animals with modified claws on only one side of the hind legs. Two of them show a supernu- merary spur (on the anterior claw in one VOLUME 111, NUMBER 3 667 Table 1.—Measurements and pt index of the holotype* (786.1 jm in length) and nine paratypes of M. nelsonae sp. n. al btl btd ssi prl 1°pl 2°pl ecl1° icl1° ecl3° ic]3° pel4° acl4° ym embryo 42.6 Se) B47 16.8 9.9 5.0 pt 13.8 81.5 39.4 23.2 13.8 pm 230.0 33.7) 5.0 27.7 149 7.9 5.9 7.9 6.9 8.9 VD 9.9 8.9 pt 148 82.2 442 23.4 17.5 23.4 205 264 23.4 294 26.4 pm 258.7 34.0 40 27.0 16.8 8.9 59 8.9 7.9 8.9 7.9 9.9 8.9 pt 118 79.4 494 26.2 Neb BOP B37) AS 8) PDI DO pm 437.8 46.5 6.9 37.6 26.7 149 10.9 10.9 BO Wl) 99 IAM Ws pt abs} | GhOK) Ab BL) Bab WBE 3) DDS Mil W/o SS pm 587.1 61.4 10.9 51.5 38.6 21.8 13.9 12.9 11.9 14.9 11.9 16.8 14.9 pt 7 S839 CO 35.5) 2o Ald IAb 243) 19.4 27.4 24.3 ym 626.9 57.4 109 485 366 21.8 2 IAQ WO WS. 12.9 17.8 16.8 pt SOSA om OSeSin S810) 222-5 2255 ZONA eon | 922-50 Se) 2.973 em 786.1* 69.3 ISO Heb 4S) DWY.7/ 16.8 14.9 13.9 17.8 15.8 20.8 16.8 pt 20.1 yh) WAS AE) DAD Dil) 2X0), Mil Ags) SOHO) PAL pm 786.6 69.3 15.8 57.4 50.5 29.7 18.8 17.8 15.8 18.8 17.8 21.8 18.8 pt 22.8 82.8 69.3 42.9 27.1 DS BA Bol Zast- Bles) Boll wm 815.9 TODS eIS28) 637453) 2 4/) 18.8 16.8 15.8 16.8 15.8 19.8 18.8 pt 20/8 2) anne O29) ee XO 2B 20.7 26 24.7 em 925.4 V2 149 644 54.5 31.7 20.8 17.8 15.8 18.8 16.8 19.8 19.8 pt IDS O45 WOO ili Boo Bil As Aho Dilts AsO Ado al = animal length; btl = buccal tube length; btd = inner diameter of the buccal tube; ssi = stylet supports insertion level on the buccal tube; prl = macroplacoids row length; 1°pl = first macroplacoid length; 2°pl = second macroplacoid length; ecl = external claw length; icl = internal claw length (1° = front leg, 3° = second or third pair of legs); pcl = posterior claw length; acl = anterior claw length (4°= hind legs). case, in the posterior one in another one); another one has a third branch on a poste- rior claw. Measurements and pt indices (percentage ratio between the length of the considered structure and the length of the buccal tube; Pilato 1981) of the holotype and nine para- types are reported in Table 1. Midgut of the largest animals containing mastax of rotifers and sclerified parts of the bucco-pharyngeal apparatus of eutardi- grades. In the testis of male specimens stained with acetic carmine, spermatozoa with a particularly elongated and coiled head are visible. Large eggs (diameter from 84.7 to 129.5 4m without processes; Figs. 1G, 3). Conical processes (height from 20.8 to 31.7 wm, di- ameter of the base from 20.0 to 34.7 wm) showing a reticular pattern with meshes of very irregular, variable shape and size (Figs. 1H, 4, 5). By SEM their surfaces ap- pear annulated (Fig. 5). Top of the process- es rounded and usually without reticulation but with a fine annulation, sometimes light- ly notched. Each process is separated from the others by a double series of 11—12 po- lygonal areolations (S—6 wm in diameter) inside which no sculpture is visible (Figs. 4, 5). Remarks.—Comparisons with the type material of Macrobiotus pallarii and Ma- crobiotus aviglianae confirmed the synon- ymy between these two species as pointed out by Pilato & Binda (1977), and indicated the close relationship between Macrobiotus nelsonae and M. pallarii. In M. nelsonae the average size of the animals is decidedly larger, there is a more evident microplacoid, a more complex buccal armature with a larger posterior band of teeth, a wider buc- cal tube, larger claws and a more posterior insertion of the stylet supports on the buccal tube. The most evident difference between M. nelsonae and M. pallarii is in the eggs. 668 The processes are more than twenty per hemisphere in M. pallarii (checked on type material) and always less than fifteen in M. nelsonae; they are also much longer. The areolations at the base of the processes are fewer in number in M. pallarii (8—9 instead of 11-12 in M. nelsonae) and never in a double series. In M. nelsonae the pt index seems to change with the size of the animal. Comparative studies on other species may provide further informations. Repositories.—Holotype, 88 paratypes and 54 eggs are in R. Bertolani’s collection in the Department of Animal Biology at the University of Modena; 473 paratypes and about 260 eggs in D. R. Nelson’s collection in the Department of Biological Sciences of East Tennessee State University; 5 para- types and 6 eggs are in the National Mu- seum of Natural History, Smithsonian In- stitution of Washington, D.C. Murrayon stellatus, new species (Figs. 6-8, Table 2) Etymology.—From stellatus (Latin), meaning “‘starry,”’ for the star-like dots on the cuticle of the animals. Paratypes.—6 paratypes mounted in polyvinyl-lactophenol and Hoyer’s medium and 1 egg mounted in polyvinyl-lactophen- ol. Holotype.—Length 234.1 «wm, collected in January 1996 and mounted in Hoyer’s medium, slide labelled as 5NO8b-S12. Type locality.—Roan Mountain (Carter County, Tennessee, U.S.A.), on the north- facing slope at elevations of 1500 m asl in leaf litter from beech trees. Diagnosis.—Cuticle punctated, with ev- ident dorsolateral bands of relatively large star-like dots. Three rod-shaped macropla- coids, microplacoid absent. Eggs laid free- ly, with small rod-shaped processes. Description.—From 162.1 to 234.1 pm in length, colorless, without eye-spots. En- tire cuticle has a fine punctuation; more- over, two dorsolateral bands (width about 10 pm and visible at low magnification, 5 X PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON objective) of star-like dots are also very vis- ible (Figs. 6A, 7). They run through each side of the body and join anteriorly at the front of the animal and posteriorly just in front of the hind legs. The stars seem finer and closer together on the front of the ani- mal and in sections above each leg, where they cover the basal portion of the dorsal surface (Fig. 7). Peribuccal lamellae evi- dent. In these specimens the components of the buccal armature are not identifiable. Stylet supports inserted on the buccal tube at about two-thirds of its length. Shape of the strengthening bar characteristic of the genus, with a showy ventral hook. Buccal tube relatively large. Pharyngeal bulb round (ratio length/width 1.1—1.2:1), containing large apophyses and three slender rod- shaped macroplacoids (Fig. 6B). The first is the longest; the second, very close to the first, the shortest; the third shows an evident subterminal constriction. Legs long. Claws small, of ‘pullari’ type (V-shaped and with a peduncle, quadrate in lateral view; Figs. 6C, D), larger on the hind legs. Main branch of the claws with evident accessory points. Small smooth lunulae at the base of each claw. Measurements and pt indices of the ho- lotype and two paratypes, compared with two paratypes of Macrobiotus dianeae and two specimens of Macrobiotus hibernicus (coll. Kristensen), are reported in Table 2. A small broken egg was found at the type locality and attributed to this species be- cause of its similarity with that of the re- lated species M. dianeae and M. hibernicus. The egg has rod-shaped processes of 3—4 11m in height with a large base and a small- er head shaped like that of a nail (Fig. 6E). The processes look often curved. Some pro- cesses have a short thin spine on their heads. At some points a hyaline matrix cov- ering the processes is visible. The processes are aligned to include wide polygonal areas, sometimes incomplete. The surfaces of these areas seem smooth. Remarks.—Murrayon stellatus is similar to Macrobiotus dianeae and to Macrobiotus VOLUME 111, NUMBER 3 669 A Ge CA Fig. 6. Holotype of Murrayon stellatus sp. n. A, Habitus (lateral view); B, Bucco-pharyngeal apparatus (lateral view); C, Claws of the third pair of legs; D, Claws of the hind legs; E, Egg detail. Note in B the hook on the ventral tube (arrow) and in C and D the quadrate peduncle at the bases of the claws (asterisks). Scale bars, 20 um for A; 10 wm for B, E; 5 wm for C, D. 670 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Measurements and pt index of the holotype* (234.1 tm in length) and two paratypes of M. stellatus sp. n., two specimens of M. hibernicus and two paratypes of M. dianeae. al btl btd ssi pri 1°pl 2°pl 3°pl ecl1° icl1° ecl3° ic]3° pel4° acl4° M. stellatus sp. n. pm 214.1 29.7 40 20.8 12.9 5.0 3.0 40 — — 5.9 5.9 6.9 6.9 pt 13.5 70.00 43.4 16.8 10.1 13.5 — — 19:9" 1939232 e2s pm NOM aed, 40 20.8 — — — — 5.9 5.9 S8) 5.9 6.9 6.9 pt 14.4 75.1 — — — = 21:3 21.3. 213, 23 e242! pm 234.1* 29.7 3.00 20.8 13.9 5.0 3.0 4.0 5.9 5.9 5.9 5.9 6.9 6.9 pt 10.1 70.00 468 168 101 135 199 199 199 199 23.2 23.2 M. hibernicus pm 204.0 23.8 2.00 15.8 10.9 4.0 3.0 4.0 4.0 4.0 4.0 4.0 4.0 4.0 pt 84 664 45.8 168 126 168 168 168 168 16.8 16.8 16.8 pm 23902 2.0 188 LES 4.0 3.0 4.0 5.0 5.0 5.0 5.0 5.0 5.0 pt nS “13.2 463° 1516 tie7, 156) 195) 195 9s aS See SS M. dianeae wm 204.0 24.8 ZO G: Se eS) 4.0 3.0 4.0 4.0 4.0 4.0 4.0 — — pt Sal 6767 AS01612 G1 Gs Gr Gs aGe — — em 259.0 31.7 3.0 23.8 14.9 5.0 3.0 5.0 6.9 6.9 6.9 6.9 6.9 6.9 pt 95 75.1 47.0 15.8 9 15:8 21:8 ~ 21:8) 218) 21 See Seeeeles al = animal length; btl = buccal tube length; btd = inner diameter of the buccal tube; ssi = stylet supports insertion level on the buccal tube; prl = macroplacoid row length; 1°pl = first macroplacoid length; 2°pl = second macroplacoid length; 3°pl = third macroplacoid; ecl = external claw length; icl = internal claw length (1° = front leg, 3° = second or third pair of legs); pcl = posterior claw length; acl = anterior claw length (4° = hind legs). hibernicus. The original description of M. hibernicus was based on an animal artifi- cially extracted from the egg (Murray 1911). Murray described three macropla- coids of similar size, twice as long as wide, and a microplacoid, but he did not cite any cuticular sculpture. This structure was re- ported by Cuénot (1932), who described a punctated band of very fine grains that forms a connecting band or bridge on the dorsal surface (at a level corresponding to the pharyngeal bulb) and that continues down on the sides of the animal, running longitudinally to the caudal extremity. Cuénot added that the second macroplacoid is a little shorter than the first one and in contact with it, the third is the longest, and the microplacoid is present, but can also be absent. The attribution of the Cuénot’s ma- terial to M. hibernicus is not certain, but very probable because of the considerable similarity of the eggs with that described by Murray (1911). The same aspect of the cu- ticular sculpture described by Cuénot is present in specimens of M. hibernicus from Val Piantonetto, Piemonte, Italy, in Mauc- ci’s collection and from Godhavn, Green- land, in Kristensen’s collection. With light microscopy M. stellatus, M. hibernicus, and M. dianeae show a similar type of cuticle. The cuticle of M. dianeae was studied by TEM and defined as a “‘heterotardigrade type’, with a mucous layer, outer laminary layer, epicuticle separated in two layers by means of pillars (rods), inner trilaminary layer and procuticle’’ (Kristensen 1982). Moreover, these species have very similar claws and bucco-pharyngeal apparati (see following sections: further systematic re- sults and discussion). Murrayon stellatus differs from M. dianeae in having a more evident cuticular sculpture, very visible even with low magnification, in having a clearly rounded bulb and in having the first macroplacoid longer than the other two. Moreover the claws of M. dianeae are de- VOLUME 111, NUMBER 3 671 Figs. 7-8. Murrayon stellatus sp. n. 7, Sculpture of the dorsolateral cuticle (DIC); 8, Cuticle with pillars (arrow; DIC). Scale bars, 1 wm. cidedly smaller in animals of similar length. M. stellatus differs from M. hibernicus in the type of the punctation of the cuticle, which in M. hibernicus is hardly visible and only at high magnification (although dor- solateral bands are present in some ob- served specimens but clearly less evident than in M. stellatus), in having a larger buc- cal tube and the hind claws longer than the first three pairs. Also the egg is very similar to those of M. hibernicus and M. dianeae, but it differs from both in that the heads of the processes are smaller in diameter and also short spines are present on the pro- cesses of M. stellatus but absent in M. hib- ernicus and longer and very visible in M. dianeae. Repositories.—Holotype, 4 paratypes and an egg are in R. Bertolani’s collection in the Department of Animal Biology at the Uni- versity of Modena, 2 paratypes are in D. R. Nelson’s collection in the Department of | Biological Sciences at East Tennessee State University; a paratype is in the National Museum of Natural History, Smithsonian Institution of Washington, D.C. Further Systematic Results Light microscopy observations on spec- imens of Macrobiotus hibernicus and Ma- crobiotus dianeae revealed that these two species are characterized by claws of the ‘pullari’ type, i.e., V-shaped with the two branches that diverge from one another at the base, forming an acute angle, and with the typical quadrate peduncle at the base (Bertolani & Pilato 1988). Both also have an evident hook on the ventral margin of the strengthening bar of the buccal tube. A personal communication from R. M. Kristensen on the presence of pillars in the cuticle of Murrayon hastatus suggests the 672 need for further investigations on the cuti- cle of the genus. Discussion The morphological observations on Mur- rayon pullari, Murrayon hastatus, Macro- biotus hibernicus and Macrobiotus dianeae allow reanalysis of the systematic position of some of these species and the opportu- nity to define the characteristics of the ge- nus Murrayon. Based on the observations of abundant material of M. dianeae (where claws were very visible in profile), this spe- cies had claws of the ‘pullari’ type, and not of the ‘hufelandi’ type with a short basal tract, as reported by Bertolani & Pilato (1988). The presence of ‘pullari’ type claws and of a ventral hook on the strengthening bar are evidence that M. hibernicus and M. di- aneae should be transferred to the genus Murrayon. The genus Murrayon is there- fore composed of the following 7 species: M. pullari (type species), M. hastatus, Mur- rayon hibernicus comb. n., Murrayon no- centiniae (Ramazzotti 1961), Murrayon di- aneae comb. n., Murrayon ovoglabellus (Biserov 1988) and Murrayon stellatus. The discovery of M. stellatus and the transfer of M. hibernicus comb. n. and M. dianeae comb. n. indicate that the genus Murrayon is not exclusively limnic, as it was previously considered. Systematic Key to the Genus Murrayon 1. Cuticle superficially sculptured with more or less evident punctation (dots) 2 — Cuticle superficially smooth, without punctation 2. Cuticle uniformly and finely punctated, egg processes immersed in a hyaline ma- trix, bearing spines and delimiting po- lygonal areas M. dianeae — Cuticle with two dorsolateral bands of more evident dots 3. Dorsolateral bands of dots very evident, egg processes with small heads, some bearing short spines on the top ....... PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON — Dorsolateral bands of dots barely evi- dent, egg processes without any spine M. hibernicus 4. Long thin macroplacoids, the first with a deep constriction, the second with an ev- ident subterminal narrowing, egg with large processes covered by a hyaline ma- trix M. hastatus — Short wide macroplacoids 5. First macroplacoid with a slight median constriction, smooth egg .. M. ovoglabellus — First macroplacoid very constricted in the middle; eggs with processes separat- ed from one to another and inserted on a smooth shell ..)5 04: 932s ae 6 6. Eggs with small, rigid, conical tubercles, with surface of the egg visible M. pullari — Eggs with long conical and bent pro- cesses with aspect of large aculeus .... M. nocentiniae Acknowledgements I would like to thank L. Rebecchi (Uni- versity of Modena, Italy) for giving me her suggestions; R. M. Kristensen (University of Copenhagen, Denmark) for his sugges- tions; R. M. Kristensen and G. Pilato (Uni- versity of Catania, Italy) for sending me samples; C. Morlini and D. Giannetti (Uni- versity of Modena) for helping me with the drawings; the Museum of Natural History of Verona (Italy) for giving me the oppor- tunity to examine Maucci’s collections. Special thanks to R. Bertolani (University of Modena, Italy), D. R. Nelson (East Ten- nessee State University, Johnson City, U.S.A.) and their collaborators for their help and suggestions. This study was sup- ported by a grant from the University of Modena for a foreign countries fellowship and by a MURST 60% grant. Literature Cited Bertolani, R. 1981. The taxonomic position of some eutardigrades.—Bollettino di Zoologia 48:197— 203. , & R. M. Kristensen. 1987. New records of Eohypsibius nadjae Kristensen, 1982, and re- vision of the taxonomic position of two genera of Eutardigrada (Tardigrada). Pp. 359-372 inR. VOLUME 111, NUMBER 3 Bertolani, ed., Biology of Tardigrades. Selected Symposia and Monographs U.Z.I., vol. 1. Muc- chi, Modena, Italy. , & G. Pilato. 1988. Struttura delle unghie nei Macrobiotidae e descrizione di Murrayon n. gen. (Eutardigrada)—Animalia 15:17—24. , & L. Rebecchi. 1993. A revision of the Ma- crobiotus hufelandi group (Tardigrada, Macro- biotidae), with some observations on the taxo- nomic characters of eutardigrades.—Zoologica Scripta 22:127—152. Biserov, V. I. 1990a. On the revision of the genus Ma- crobiotus. The subgenus Microbiotus s.str.: a new systematic status of the group hufelandi (Tardigrada, Macrobiotidae). Communication 1.—[In Russian] Zoologicheskii Zhurnal 69:5— 17. . 1990b. On the revision of the genus Macro- biotus. The subgenus Macrobiotus s.str. is a new taxonomic status of the hufelandi group (Tardigrada, Macrobiotidae). Communication 2.—[In Russian] Zoologicheskii Zhurnal 69:37— 50. Cuénot, L. 1932. Tardigrades. pp. 96. Jn Faune de France. vol. 24. ed. Paul Lechevalier, Paris. Guidi, A., & L. Rebecchi. 1996. Spermatozoan mor- phology as a character for tardigrades system- atic: comparison with sclerified parts of animals and eggs in eutardigrades.—Zoological Journal of the Linnean Society 116:101-113. Kristensen, R. M. 1982. New aberrant eutardigrades from homothermic springs on Disko Island, 673 West Greenland. Pp. 203—220 in D. R. Nelson, ed., Proceedings of the Third International Sym- posium on the Tardigrada. East Tennessee State University Press, Johnson City, Tennessee. Marcus, E. 1936. Tardigrada. Pp. 340 Jn Das Tierreich, vol 66. Walter de Gruyter, Berlin und Leipzig. Murray, J. 1911. Arctiscoida. Proceedings of the Royal Irish Academy 37:1—16. Pilato, G. 1969a. Evoluzione e nuova sistemazione de- gli Eutardigrada.—Bollettino di Zoologia 36: 327-345. . 1969b. Schema per una nuova sistemazione delle famiglie e dei generi degli Eutardigrada.— Bollettino dell’ Accademia Gioenia di Scienze Naturali, Catania Ser. 4% 10:181—193. . 1981. Analisi di nuovi caratteri nello studio degli eutardigradiAnimalia 8:51—57. Pilato, G., & M. G. Binda. 1977. Precisazioni e retti- fiche alla descrizione di alcune specie di tardi- gradi (seconda nota).—Animalia 4:35-51. Ramazzotti, G. 1962. If Phylum Tardigrada—Memo- rie dell’Istituto Italiano di Idrobiologia 16:1— S25). . 1965. II Phylum Tardigrada (1° suppl.).—Me- morie dell’Istituto Italiano di Idrobiologia 19: 101-212. Schuster, R. O., D. R. Nelson, A. A. Grigarick, & D. Christenberry. 1980. Systematic criteria of the Eutardigrada.—Transactions of the American Microscopical Society 99:284-303. Thulin, G. 1928. Ueber die Phylogenie und das System der Tardigraden.—Hereditas 11:207—266. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):674—693. 1998. The genus Perinereis (Polychaeta: Nereididae) from Mexican littoral waters, including the description of three new species and the redescriptions of P. anderssoni and P. elenacasoae Jests Angel de Leén Gonzalez and Vivianne Solis-Weiss (JALG) Divisi6n de Biologia Marina, Centro de investigaciones Biolédgicas del Noroeste, S.C., Ap. Postal 128. La Paz, B.C.S. 23000, Mexico; (VS-W) Laboratorio de Ecologia Costera, Instituto de Ciencias del Mar y Limnologia, U.N.A.M.., Ap. Postal 70-305, Mexico, D.E 04510, Mexico Abstract.—Eight species of the genus Perinereis were found in Mexican waters (Gulf of Mexico, Caribbean Sea and Pacific Ocean). Of these, three species are newly described: one for the Caribbean (P. cariboea) and two for the Mexican Pacific (P. bajacalifornica and P. osoriotafalli). Perinereis an- derssoni is redescribed based on type material collected in Rio de Janeiro, Brazil. The previous records of this species from Mexico are referred to P. elenacasoae. A neotype is designated for P. elenacasoae, since the type ma- terial is lost, and the species is redescribed based on the neotype. The genus Perinereis is characterized by the presence of conical paragnaths on both pharyngeal rings of the proboscis, except in Area VI, where either transverse long rib- bon-shaped, short transverse bars, or a row of small bars occur. Bars may be so short as to appear cone-like on some species (Hutchings et al. 1991). Notopodia can be enlarged posteriorly or not modified. Only compound homogomph spinigers present in notopodia. Homogomph and heterogomph spinigers and heterogomph falcigers are present in neuropodia. The terminology of parapodial structures was taken from Fig. 1 of Hutchings & Reid (1990). Previously, four species in this genus have been reported from littoral waters of Mexico: P. anderssoni Kinberg, 1866, P. elenacasoae Rioja, 1947, P. monterea (Chamberlin, 1918) and P. villalobosi Rio- ja, 1947. However, the specimens so far re- corded as P. anderssoni from mexican shores actually belong to P. elenacasoae. This confusion arose from a description and illustrations by Rioja (1960: 296) of speci- mens from the Mexican region of the Gulf of Mexico incorrectly reported as P. an- derssoni. For this reason, a redescription of P. anderssoni is undertaken, based on type material reported by Kinberg (1866) from Rio de Janeiro, Brazil. Perinereis elenaca- soae is redescribed based on material from the type locality: Mazatlan, Sinaloa, and P. villalobosi is additionally described based on specimens from La Paz, Baja California Sur. Perinereis obfuscata Grube, 1878 was referred to P. elenacasoae by Salazar Val- lejo (1989), based on specimens collected in western Mexico. This synonymy was found to be correct when the material de- posited by Berkeley & Berkeley (1960) in the Smithsonian Institution collections (USNM) from Zihuatanejo coasts was ex- amined by one of us (J.A.L.G). For this study, specimens in the collec- tions of the Natural History Museum of Los Angeles County (LACM-AHP), U.S. Na- tional Museum of Natural History, Smith- sonian Institution (USNM), Naturhistoriska Rijksmuseet Stockholm (NRS), the Instituto de Ciencias del Mar y Limnologia, UNAM (CPICML) and the Universidad Aut6noma de Nuevo Leon (UANL) were examined. VOLUME 111, NUMBER 3 Genus Perinereis Kinberg, 1866 Perinereis anderssoni Kinberg, 1866 Fig. 1A-—G Perinereis anderssoni Kinberg, 1866: 175.—Hartman, 1951:47, pl. 13, Fig. 6.—Fauchald, 1977:31, Fig. 8a—b. Material examined.—Type series con- sisting of 13 poorly preserved specimens collected in Rio de Janeiro, Brazil (NRS- 156); other material: Punta da Cruz, Ilha de Sao Francisco, Santa Catharina, Brazil, 28 Oct 1925 (1 specimen) (USNM 24229); Juan Fernandez, Chile, 8 Dec 1926 (7 spec- imens) (USNM 24252). Campeche: Puerto Real, Ciudad del Carmen, Mexico, M. E. Caso, coll., 31 Jul 1972 (1 specimen). Redescription.—Best preserved speci- men of type series complete with 85 setig- ers; 63 mm long, 4 mm wide, including par- apodia, with no evident pigmentation. Prostomium longer than wide, two pairs of eyes in trapezial arrangement. Frontal antennae short. Biarticulate palps large with conical palpostyle. Peristomium with four pairs of short tentacular cirri, longest pair extending posteriorly past first setiger (Fig. 1A). Paragnaths of the pharyngeal areas ar- ranged as follows: I, 4 cones in triangle; II, 13 cones in a triangle; III, 19 cones in 3 rows; IV, curved group of 27 cones; V, 3 cones in triangle; VI, short transverse bar; VII-VIII, 45 cones in 2 rows. Jaws with 5 teeth (Fig. 1B). First two parapodia uniramous, dorsal and ventral cirri subequal (Fig. 1C). Fol- lowing anterior parapodia biramous with dorsal and median notopodial ligules ante- riorly rounded; neuropodial postsetal lobes truncate, superior lobe conical, inferior one rounded; ventral neuropodial ligule slender. Dorsal cirri proximally inserted. Median parapodia with dorsal and median notopo- dial lobes conical (Fig. 1D), neuropodial structures similar to those in anterior para- podia; dorsal cirri medially inserted. Pos- terior parapodia with dorsal cirri inserted medially and anteriorly, notopodial ligule 675 enlarged; median notopodial ligule subu- late; postsetal neuropodial lobes mamilli- form, ventral neuropodial ligules rounded anteriorly; ventral cirri digitiform and in- serted proximally (Fig. 1E). Uniramous parapodia with supracicular homogomph spinigers, and infracicular het- erogomph falcigers. Biramous parapodia with setation similar throughout body. No- topodial supracicular setae homogomph spinigers. Neuropodial supracicular setae homogomph spinigers and heterogomph falcigers; infracicular neurosetae hetero- gomph spinigers and falcigers. Infracicular neuropodial falcigers in anterior parapodia with distally blunt appendage (Fig. 1F); ap- pendage in posterior parapodia slender and distally pointed (Fig. 1G). Pygidium with terminal anus, with pair of short ventrally inserted anal cirri. Distribution.—Amphi-American. Origi- nally described from Rio de Janeiro, Brazil. The species is distributed in the Atlantic and Gulf of Mexico (Ciudad del Carmen, Campeche, Mexico), eastern Gulf of Mex- ico (Hartman, 1951) and in the Pacific, in Juan Fernandez Island (Chile). Habitat.—Not defined in records. Perinereis bajacalifornica, new species Fig. 2A—F Material examined.—Baja California Sur: Falsa Bay, La Paz, W. H. Shepherd, coll., 22 Sep 1971, (1 specimen) (USNM 48858); Balandra mangrove, La Paz, E. Amador coll., 24 Sep 1985, (1 specimen, Holotype) (USNM 180693); Zacatecas mangrove, La Paz, same collector, 12 Feb 1986 (8 specimens). Description.—Holotype complete with 93 setigers, 40 mm long and 2.5 mm wide including parapodia. Body pale yellow, with no evident pigmentation pattern. Prostomium pentagonal, with two pairs of eyes, anterior ones oval and slightly more separated from each other, posterior ones rounded. Pair of short cirriform anten- nae. Biarticulate palps with conical palpo- 676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Perinereis anderssoni. A. Anterior region dorsal view; B. Everted pharynx, dorsal view from a specimen in the type group; C. Ist parapodium; D. Parapodium 39th; E. Parapodium 71th; K Heterogomph neuropodial falciger in infracicular position from setiger 10; G. Same of setiger 65. (Scale: A, B = 1 mm; C, D, E = 100um; EF G = 15pm). styles. Peristomium with longest pair of naths; II, 11 conical paragnaths arranged in tentacular cirri extending to setiger 3 (Fig. triangle; III, 15 cones in trapezial group; IV, 2A). curved group of 21 cones; V, single cone; VI, Paragnaths arranged on pharyngeal areas transverse, slender, ribbon-shaped bar; VU-— as follows: I, a clump of 7 conical parag- VIII, row of 7 small cones. (Fig. 2B, C). VOLUME 111, NUMBER 3 Anterior parapodia with notopodial and neuropodial lobes and ligules conical, subequal; dorsal cirri stout not longer than dorsal ligules (Fig. 2D). Median parapo- dia with notopodial dorsal ligules slightly longer than other lobes; dorsal cirri simi- lar to those of anterior parapodia and proximally inserted. Posterior parapodia with notopodial dorsal ligules conspicu- ously longer, distally conical and dorsally pigmented; dorsal cirri slender, medially inserted, not longer than the dorsal lig- ules. Notopodial median ligules conical. Neuropodial postsetal lobes rounded, ven- tral ligules digitiform, slightly longer than neuropodial postsetal lobes. Ventral cirri slender (Fig. 2E). Supracicular homogomph spinigers in notopodia. Supracicular homogomph spi- nigers and heterogomph falcigers in neu- ropodia, but only homogomph spinigers in anterior parapodia. Neuropodial infracicular setae consisting of single homogomph spi- niger and some hetrogomph falcigers; latter with 9-10 teeth on margin (Fig. 2F). Pygidium with two short slender anal cir- ri; anus terminal. Discussion.—Perinereis bajacalifornica, n. sp., belongs to the group of species in which a ribbon-shaped transverse bar is present in Area VI of the proboscis and no- topodial lobes are enlarged on posterior se- tigers. In their revision, Hutchings et al. (1991) included 16 species in this group: P. amblyodonta (Schmarda 1861) and P. bar- bara (Monro 1926) from Australia, P. an- derssoni Kinberg, 1866 and P. pontoni Kin- berg, 1866 from Brazil, P. elenacasoae Rioja, 1947, from western Mexico, P. falk- landica Ramsay, 1914, for the Falkland is- lands, P. longidonta Rozbaczylo & Castilla, 1973 from Chile, P. macropus (Claparéde 1870) from the Mediterranean Sea, P. ma- layana (Horst 1889), P. nigropunctata (Horst 1889) and P. tobeloana (Augener 1933) from the Malay Archipielago, P. obfuscata (Grube 1878) and P. sululana (Horst 1924) from the Philippines, and P. pseudocavifrons Fauvel, 1930 from New 677 Caledonia. Perinereis bajacalifornica, n. sp., differs from the other species in this group by having a single row of seven small cones on pharyngeal areas VII—VIII. In the other species of the group, a variable number of conical paragnaths, arranged in 2—4 rows and varying in numbers from 18 to 101, is present. Perinereis elenacasoae, described from western Mexico, is biogeographically the most closely related species to P. bajacal- ifornica, n. sp. These species can be differ- enciated from each other and from other species in the group mainly by the arrange- ment of their paragnaths. Etymology.—The specific name is de- rived from the name of the State of Baja California Sur, Mexico. Distribution.—This species is only known from the mangrove areas in La Paz, Baja California Sur, Mexico. Habitat.—In sediment trapped among mangroves. Perinereis cariboea, new species Fig. 3A—-E Material examined.—Quintana Roo: As- cencion Bay, Punta Pajaros, V. Solis-Weiss, coll. 6 Oct 1983 (Holotype, USNM 180694). Yucatan: Cerritos Island, S. I. Sa- lazar Vallejo, coll. 18 Jan 1991 (6 speci- mens). Description.—Holotype complete with 53 setigers, 35 mm long, 1.5 mm wide, green with strongly pigmented prostomium. Pharynx not everted in specimens. Anal- ysis performed by dissection. Prostomium pentagonal, with dorsal groove, with two pairs of rounded eyes in quadrate arrangement. Pair of short digiti- form frontal antennae. Biarticulate palps globose, with small, conical palpostyles. Peristomium with four pairs of tentacular cirri, anterior pair extending posteriorly to setiger 4 (Fig. 3A). Paragnaths arranged on pharyngeal areas as follows: I, 2 cones; II, 8 cones in 2 rows; III, oval group of 7 cones; IV, triangular 678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. in ventral view; D. Parapodium 10th; E. Parapodium 80th; E Heterogomph neuropodial falciger in supracicular position from setiger 80. (Scale: A, B, C = 0.5 mm; D, E = 150m; F = 10pm). group of 12 cones; V, no paragnaths; VI, 2 transverse flattened bars, VII-VIII, 11 cones in 2 rows. Anterior notopodia with dorsal and ven- tral ligules conical, neuropodial postsetal lobes rounded, ventral ligules enlarged; dorsal cirri inserted proximally, stouter than the ventral cirri (Fig. 3B). Median parapo- Perinereis bajacalifornica, n. sp. A. Anterior end, dorsal view; B. Pharynx in dorsal view; C. Pharynx dia with notopodial dorsal ligules proxi- mally enlarged, median ligules, neuropodial postsetal lobes and ventral ligules rounded; dorsal cirri inserted medially (Fig. 3C). Posterior parapodia with notopodial dorsal ligules enlarged; median ligules, neuropo- dial postsetal lobes and ventral ligules sim- ilar to those of median parapodia. Dorsal VOLUME 111, NUMBER 3 679 Fig. 3. Perinereis cariboea, n. sp. A. Anterior end, dorsal view; B. Parapodium 10th; C. Parapodium 30th; D. Parapodium 50th; E. Heterogomph neuropodial falciger in supracicular position from setiger 30. (Scale: A = 0.5 mm; B, C, D = 100m; E = 8pm). cirri inserted near distal margin of dorsal ligules (Fig. 3D). Setation similar throughout body as fol- lows: notosetae all homogomph spinigers; supracicular neurosetae homogomph spini- gers and heterogomph falcigers; infracicu- lar neurosetae heterogomph, represented by single spiniger and 4 falcigers (Fig. 3E). Pygidium with a pair of short anal cirri; anus termino-dorsal. Discussion.—Perinereis cariboea, 0. sp., belongs to the group of species in which 680 two transverse flattened bars are present in area VI of the proboscis and notopodial dorsal ligules are enlarged posteriorly. Only two other species are known in this group: P. mochimaensis Lifiero-Arana, 1983 de- scribed from Venezuela, and P. osoriota- falli, n. sp, herein described. These species can be differentiated from each other by the ornamentation of pharyngeal areas I, V and VII-VIII. In P. cariboea, n. sp., two small cones in a row are present on area I, area V lacks paragnaths, and 11 cones in two rows are present on areas VII—VIII. In P. mochimaensis, 11 cones in a triangle are present on area I, two cones are present on area V and 32 cones in two rows are present on areas VII—VIII. In P. osoriotafalli, n. sp., four cones are present on area I in trapezial arrangement, one cone is present on area V, and eight cones in a row are present on ar- eas VII-VIII. Etymology.—The specific name is de- rived from the name of the Caribbean Sea. Distribution.—Mexican Caribbean; only known from two localities: Concepcion Bay, Quintana Roo State and Cerritos Is- land, Yucatan. Habitat.—Among algae attached to rocks in the intertidal zone. Perinereis elenacasoae Rioja, 1947 Figs. 4A—E, 5A—D Perinereis elenacasoi Rioja, 1947: 531, Figs. 8-17. Perinereis elenacasoae Salazar Vallejo, 1989:50 Perinereis obfuscata Berkeley & Berkeley, 1960:359 Perinereis anderssoni Rioja, 1960:295. Figs. 12-15. Material examined.—Baja California Sur: La Paz Bay, W. M. Shepherd, coll., Aug 1976 (10 specimens); Caimancito beach, La Paz Bay, J. A. de Le6n-Gonzalez, coll., 12 Oct 1987 (2 specimens). Sonora: Penasco Harbor, in front of shrimp farm, 5 May 1981, same collector (2 specimens); same locality, M. Silva, coll., 7 Jan 1985 (1 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON specimen); same locality, Norse beach, V. A. Gallardo, coll. no date (6 specimens). Sinaloa: Mazatlan, Est. 14309, Reef of southern lighthouse island, E. Y. Dawson, coll., 12 Jul 1946 (1 specimen); same lo- cality and collector, Est. 14308, north Olas Atlas lighthouse, 12 Dec 1946 (3 speci- mens); same locality and collector, Est. 14311, 2 miles north of Mazatlan, 6 Jul 1952 (1 specimen); same locality, Cerritos beach, J. A. de Leon-Gonzalez, coll., 15 May 1981 (Neotype, USNM 180695) and 6 specimens; Mazatlan area, collector un- known, 1986 (8 specimens); same locality, Casa del Marino, collector unknown, 1986 (16 specimens); same locality, Chivos Is- land, collector unknown, 1986 (10 speci- mens). Nayarit: Larga Island, Marietas Is- lands, C. Lépez Rivas, coll., 17 Dec 1994 (1); same, 13 Feb 1995 (5 specimens). Jal- isco: Barra de Navidad, Est. 14310, E. Y. Dawson, coll., 25 Dec 1946 (3 specimens). Guerrero: Acapulco Harbor, San Lorenzo reef, Est. 1552-46, E. Y. Dawson, coll., 9 Apr 1946, (4 specimens). Oaxaca: Salina Cruz, collector unknown, 27 May 1951 (1 specimen). Veracruz: Veracruz Harbor, Pla- ya Norte reef, E. Rioja, coll., Jan 1960 (12 specimens); Boca del Rio, north jetty, M. L. Jones, coll., 8 Aug 1962 (60 specimens); Tuxpam, north rocky area, H. Gonzalez, coll., 25 May 1978 (3 specimens); same lo- cality, G. Gongora-Garza, coll., 05 Jun 1984 (3 specimens); Barra de Cazones, Pul- po point, J. A. de Leé6n-Gonzalez, coll., 1 Jun 1985 (9 specimens); same, 15 Jun. 1985 (37 specimens); same, 31 Oct 1991 (93 specimens); Barra of Tamiahua, north rocky area, J. A. de Leén-Gonzalez, coll., 5 May 1985 (1 specimen); same locality, G. Guajardo, coll., 25 May 1991 (3 speci- mens); same, 8 Jun 1991 (4 specimens). Campeche: Puerto Real, Ciudad del Car- men, M. E. Caso, coll., 31 Jul 1972 (1 spec- imen). Quintana Roo: Smithsonian-Bredin Expedition IV: Mujeres Island, stn. 29-60, 31 Mar 1960 (1 specimen); Espiritu Santo Bay, stn. 41-60, 6 Apr 1960 (3 specimens); Allen Point, Ascencién Bay, stn. 45-60, 07 VOLUME 111, NUMBER 3 Apr 1960 (2 specimens), Nicchehabin reef, Ascencion Bay, stn. 67-60, 13 Apr 1960 (3 specimens); same, stn. 72-60, 14 Apr 1960 (1 specimen); same, stn. 91-60, 18 Apr 1960 (1 specimen); Ascencion Bay, V. So- lis-Weiss, coll., 6 Oct 1983 (1 specimen); Cozumel Island, same collector, 10 Oct 1983 (2 specimens). Epitokous material examined.—Guerre- ro: Zihuatanejo, W. L. Klawe, coll., 6 Sep 1958 (1 female), (USNM 33496). Description of the Neotype.—Specimen complete with 72 setigers, 34 mm long and 2 mm wide including parapodia; color pale yellow, vestigial dark pigmentation in an- terior region. Prostomium pentagonal, two pairs of eyes in quadrate arrangement. Frontal an- tennae stout, digitiform. Biarticulate palps slender with globose palpostyles. Peristo- mium with four pairs of relatively short ten- tacular cirri, longest pair extending poste- riorly to second setiger (Fig. 4A). Pharynx not everted in specimens. Anal- ysis performed by dissection. Pharynx with paragnaths arranged on pharyngeal areas as follows: I, group of 11 cones; II, 25 cones in trapezial arrange- ment; III, oval group of 25; IV, curved group of 16—17 cones in 4 rows; V, single cone; VI, transverse flattened, ribbon- shaped bar; VII—VIII, 37 cones in 2 rows. Anterior parapodia with notopodial dorsal and median ligules distally rounded, neuro- podial postsetal lobes less developed; dorsal and ventral cirri subequal (Fig. 4B). Median parapodia with notopodial and neuropodial lobes and ligules conical, neuropodial dorsal ligules longer (Fig. 4C); dorsal and ventral cirri similar to those of anterior segments. Posterior parapodia notopodial dorsal ligules enlarged, dorsal cirri medially inserted; me- dian ligule and neuropodial postsetal lobes distally conical, ventral ligule digitiform; ventral cirri inserted proximally, with pos- terior protusion (Fig. 4D). Setation similar throughout body. Noto- podia with homogomph spinigers. Supra- cicular neurosetae homogomph spinigers 681 and stout heterogomph falcigers with ante- rior slender tooth directed downward (Fig. 4E); infracicular neurosetae heterogomph spinigers and slender heterogomph falcigers. Pygidium with a pair of short anal cirri inserted ventrally. Anus terminal. Epitokous female.—Best preserved spec- imen with 43 setigers, 14 mm long and 2.5 mm wide without parapodia. Prostomium longer than wide with a frontal median dorsal groove. Pair of small digitiform antennae, 2 pairs of large eyes in quadrate arrangement. Peristomium with 4 pairs of tentacular cirri, longer ones extend- ing posteriorly to setiger 3 (Fig. 5A). Body divided into moderately modified anterior region and heteronereidid region; anterior region with modified, proximally stout dorsal cirri in first 5 setigers (Fig. 5B); parapodia similar to those of atokous spec- imens from setiger 6 to 17. Parapodia of heteronereidid region compressed, flattened antero-posteriorly; dorsal cirri without evi- dent crenulation. Natatory lamellae associ- ated with parapodial lobes and ligules and with proximal region of dorsal and ventral cirri (Fig. 5C, D). Normal setae replaced by natatory setae with broad, paddle-shaped appendages. Discussion.—Perinereis elenacasoae Rioja, 1947, was originally described from Mazatlan, (Sinaloa State) shores; unfortu- nately the type material, as well as all the other types described by Enrique Rioja, have been lost. Based on Article 75 of the International Code of Zoological Nomen- clature, in this paper a Neotype for this spe- cies, collected in the type locality: Maza- tlan, Sinaloa, is designated. Distribution.—Amphi-American. Mexi- can Pacific from Puerto Penasco, Sonora (Gulf of California), to Salina Cruz (Oa- xaca); Atlantic Ocean from Gulf of Mexico and Caribbean Sea south to Brazil. This is the first record for the species from the western Atlantic. Habitat.—On rocky substrates, among rhizoids of algae attached to rocks, and cor- al substrates. 682 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Perinereis elenacasoae. A. Anterior end, dorsal view; B. Parapodium 10th; C. Parapodium 31th; D. Parapodium 50th; E. Heterogomph neuropodial falciger in supracicular position from setiger 50. (Scale: A = 1 mm; B, C, D = 150m; E = 10pm). VOLUME 111, NUMBER 3 683 Fig. 5. Perinereis elenacasoae (epitoke). A. Anterior end, dorsal view (parapodia omitted); B. Parapodium 4th; C. Parapodium 30th; D. Parapodium 40th. (Scale: A = 1 mm; B, C, D = 100m). Perinereis floridana Ehlers, 1868 Material examined.—Campeche: Alacra- Figs. 6A-E, 7A—E nes reef East, V. Solis-Weiss, coll., 28 Oct Perinereis floridana Ehlers, 1868: 503.— 1990 (7 specimens); same, 23 Mar 1991 (45 Salazar Vallejo & Jimenez-Cueto, 1996— specimens); Alacranes reef West, same col- 1997: 367. lector, 23 Mar 1991 (3 specimens); Arenas 684 Cay, Southwest, same collector, 21 Mar 1991 (8 specimens); Arenas Cay North, same collector 17 Mar 1991 (4 specimens). Quintana Roo: Mujeres Island, stn. 21-60, Smithsonian-Bredin Expedition IV, 30 Mar 1960 (1 specimen); Ascenci6n Bay, stn. 67- 60, same expedition, 13 Apr 1960 (2 spec- imens). Epitokous material.—Key Largo, Flori- da, USA, J. Ross, coll., 20 Dec 1950, (1 female). Diagnosis.—Best preserved specimen in- complete, 50 mm long, 3 mm wide includ- ing parapodia, with 73 setigers. Prostomium longer than wide, two pairs of small eyes in quadrate arrangement. Frontal antennae cirriform. Palps biarticu- late, palpostyles conical. Peristomium with longest pair of tentacular cirri extending posteriorly to setiger 4 (Fig. 6A). Pharynx not everted in specimens. Anal- ysis performed by dissection. Pharynx with paragnaths arranged on pharyngeal areas as follows: I, 2 cones; II, 9 cones in 2 rows; III, oval group of 16 cones; IV, long curved group of 18; V, sin- gle cone; VI, short transverse bar; VII—VIII, 24 cones in 2 rows. First two parapodia uniramous, following ones biramous. Anterior biramous parapo- dia with notopodial dorsal ligules slender distally; notopodial median ligules distally truncate; neuropodial postsetal lobes round- ed, ventral ligules subulate. Dorsal and ven- tral cirri subequal (Fig. 6B). Median and posterior parapodia with notopodial ligules, neuropodial postsetal lobes and neuropodial ventral ligules all triangular. Notopodial dorsal ligules not enlarged; dorsal cirri in- serted medially, ventral cirri inserted prox- imally (Fig. 6C, D). Setation similar throughout body: noto- podial supracicular setae homogomph spi- nigers; neuropodial supracicular setae hom- ogomph spinigers and heterogomph falci- gers, infracicular neurosetae heterogomph spinigers and falcigers; latter with slender appendage (Fig. 6E). Epitokous female.—Specimen light yel- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON low, 32 mm long and 4 mm wide, with 97 setigers. Prostomium as long as wide, with pair of slender cirriform antennae and two pairs of large, highly modified eyes; biarticulate palps with rounded palpostyles. Peristomi- um with longest pair of tentacular cirri ex- tending to setiger 5 (Fig. 7A). Body divided into moderately modified anterior region and highly modified heter- onereidid region; first five setigers of ante- rior region with modified dorsal and ventral cirri (Fig. 7B); parapodia similar to those of atokous specimens from setigers 6 to 18 (Fig. 7C). Parapodia of heteronereidid re- gion highly modified with lamellae associ- ated with parapodial lobes, as well as with ventral and dorsal cirri; lamellae trilobed and associated with ventral cirri in median parapodia (Fig. 7D), appearing bilobed in posterior region (Fig. 7E); normal setae re- placed by natatory setae with broad paddle- shaped appendages. Distribution.—Western Atlantic: Gulf of Mexico and Caribbean Sea. Reported in Mexico from Quintana Roo. Habitat.—Associated with coral rubble. Perinereis monterea (Chamberlin, 1918) Fig. 8A—F Nereis (Neanthes) monterea Chamberlin, 1918: 474. Nereis spinifera Treadwell, 1929: 5, figs. 15-20. Perinereis monterea Berkeley & Berkeley, 1958: 403.-Hartman, 1968: 557.—Banse & Hobson, 1974: 71, Fig. 18n.—Kuden- ov, 1979: 118.—Salazar Vallejo, 1985: 108, Fig. 33 a-e. Material examined.—Baja California: Ensenada Harbor, Todos Santos Bay, S. I. Salazar Vallejo, coll., 15 Apr 1983 (6 spec- imens). Baja California Sur: Isla Margarita, stn. 159, M. Cardenas, coll., 12. Jun 1948, (3 specimens) (USNM 24727). Sonora: Puerto Pefiasco, La Cholla bay, R. Dough- erty, coll., Oct 1976 (2 specimens). Diagnosis.—Best preserved specimen VOLUME 111, NUMBER 3 685 Fig. 6. Perinereis floridana. A. Anterior end, dorsal view; B. Parapodium 10; C. Parapodium 30; D. Para- podium 70; E. Heterogomph neuropodial falciger in supracicular position from setiger 30. (Scale: A: 1 mm; B, C, D = 100yum; E = 10yum). 686 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. complete, 65 mm long and 2.5 mm wide including parapodia, with 116 setigers, reddish with densely pigmented dorsal re- gion. Prostomium pentagonal, longer than wide, with two pairs of eyes in rectangular arrangement. Frontal antennae digitiform. Biarticulate palps with globose palpostyles (Fig. 8A). Peristomium with short tentacu- lar cirri, longest pair extending posteriorly to first setiger, lower pair short and stout. Perinereis floridana (epitoke). A. Anterior end, dorsal view (parapodia omitted); B. Parapodium 3th; C. Parapodium 10th; D. Parapodium 30th; E. Parapodium 60 th. (Scale: A = 1 mm; B, C, D, E = 150m). Pharynx not everted in specimens. Anal- ysis performed by dissection. Paragnaths arranged on pharyngeal areas as follows: I, single large cone; II, oval group of 19 cones; III, oval group of 51 cones in 5 rows; IV, long curved group of 60 cones, and 2 small proximal bars; V, sin- gle large cone; VI, single short, cone- shaped bar; VII—VIII, 40 cones in 4 rows, additional row of 5 cones in midventral re- gion. VOLUME 111, NUMBER 3 Anterior parapodia with dorsal and ven- tral notopodial ligules distally rounded, su- perior lobes short, digitiform; neuropodia with postsetal lobe truncate, superior lobe short, inferior lobe quadrate, ventral ligule similar in length to notopodial dorsal ligule; dorsal cirri medially inserted, ventral cirri proximally inserted (Fig. 8B). Notopodial dorsal ligules enlarged on median and pos- terior parapodia, dorsal cirri inserted ante- riorly and medially. Neuropodial ventral ligules reduced to rounded lobes on middle segments, even more reduced posteriorly (Fig. 8C, D). Notosetae homogomph spinigers on all parapodia. Anterior neuropodia with supra- cicular neurosetae homogomph spinigers and heterogomph falcigers, latter with slen- der, distally pointed appendage (Fig. 8E); infracicular neurosetae including only het- erogomph falcigers. Median and posterior parapodia with supracicular homogomph spinigers and heterogomph falcigers; latter with anteriorly rounded appendage (Fig. 8F); infracicular neurosetae heterogomph spinigers and falcigers. Pygidium with terminal anus, pair of Slender anal cirri inserted lateral to anal opening. Distribution.—Temperate to tropical wa- ters of northeastern Pacific Ocean: Canada through State of Guerrero, Mexico. Habitat.—Reported herein from under tests of barnacle Tetraclita squamosa Dar- win, 1854 in Puerto Pefiasco, La Cholla Bay, Sonora; not defined in other records. Perinereis osoriotafalli, new species Fig. 9 A-F Material examined.—Sonora: Guaymas, San Francisco Inlet, E. Y. Dawson and E Durham, colls., 18 Mar 1946 (1 specimen); Guaymas, las playas road, in front of Las Palmas, B. Burch, coll., 13 Feb 1953 (2 specimens); Puerto Pefiasco, La Cholla Bay, stn. V-1, V. A. Gallardo, coll., (1 spec- imen). Sinaloa: Topolobampo, Los Patos Is- land, B. FEF Osorio-Tafall, coll., 15 May 687 1961 (Holotype, USNSM 180696) and 3 specimens; same, J. Reddell, coll., 24 Nov 1968 (6 specimens). Description.—Holotype 42 mm long, 5 mm wide including setae, complete with 86 setigers; color pale yellow, no evident pig- mentation pattern. Prostomium subpentagonal, longer than wide; two pairs of black rounded eyes in quadrate arrangement, anterior ones more widely separated. Pair of short digitiform antennae; palps biarticulate, palpostyles small, not everted. Peristomium with ante- rior pair of tentacular cirri, extending pos- teriorly to setiger 4, posterior pair appearing segmented proximally. (Fig. 9A). Pharynx not everted in specimens. Anal- ysis performed by dissection. Pharynx with brown, calcified jaws, with three stout teeth present on the interior mar- gin. Paragnaths arranged on pharyngeal ar- eas as follows: I, 4 cones in diamond- shaped group; II, small group of 12 cones; III, oval group of 17 cones; IV, elongate group of 23 cones; V, single cone; VI, 2 transverse, ribbon-shaped bars; VII—VIII, 8 cones in single row. Notopodial and neuropodial ligules rounded to conical on anterior parapodia; dorsal cirri digitiform, proximally enlarged, inserted on the median posterior region of notopodial dorsal ligules; ventral cirri cir- riform, proximally inserted on neuropodial ventral ligule (Fig. 9B). Notopodial dorsal ligules enlarged on median and posterior parapodia, dorsal cirri inserted on middle distal part of dorsal ligules, relatively small- er than on anterior parapodia; ventral cirri increasing considerably in size towards pos- terior end (Fig. 9C, D). Setation of anterior setigers as follows: supracicular notosetae homogomph spi- nigers, with finely serrated appendages; supracicular neurosetae homogomph spi- nigers similar to those in notopodia, and heterogomph falcigers; latter with distally rounded appendage denticulate on inner margin; infracicular neurosetae hetero- 688 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Perinereis monterea. A. Anterior end, dorsal view; B. Parapodium 9th; C. Parapodium 40th; D. Parapodium 99th; E. Heterogomph neuropodial falciger in supracicular position from setiger 9; EF Same from setiger 99. (Scale: A = 1 mm; B, C, D = 150ym; E, F = 15pm). gomph falcigers, similar to supracicular similar spinigers and falcigers (Fig. 9B), falcigers, but smaller (Fig. 9F). Median except for the presence of one pair of in- and posterior parapodia with setation sim- fracicular heterogomph neuropodial spi- ilar to that of anterior parapodia, with nigers with finely serrated appendage. VOLUME 111, NUMBER 3 689 Fig. 9. Perinereis osoriotaffali, n. sp. A. Anterior end, dorsal view; B. Parapodium 10th; C. Parapodium 40th; D. Parapodium 80th; E. Heterogomph neuropodial falciger in infracicular position from setiger 10; K Same from setiger 80. (Scale: A = 1 mm; B, C, D = 150ym; E, F = 8yum),. Pygidium with terminal anus, with pair a group that previously included only P. of short anal cirri. mochimaensis Lifiero-Arana 1983. This Discussion.—The new species is in- species group is characterized by having cluded along with P. cariboea, n. sp., in two transverse bars on Area VI and en- 690 larged notopodial dorsal ligules on pos- terior parapodia. Perinereis osoriotafalli, n. sp., can be differentiated from P. car- iboea, n. sp. and P. mochimaensis by its pharynx ornamentation, especially on area VII-VIII. Perinereis osoriotafalli has eight cones present in one row on area VII-VIII, whereas in the other two spe- cies, the cones on area VII—VIII are ar- ranged in two rows. Perinereis mochi- maensis has 32 cones on area VII—VIII, whereas in P. cariboea only 11 cones are present in this region of the proboscis. Etymology.—This species is named in honor of B. E Osorio Tafali, a close aide of Dr. E. Rioja, who collected the material upon which this species is described. Distribution.—Gulf of California: From Puerto Pefiasco to Los Patos Island, Ohuira Bay, in front of Topolobampo Harbor. Habitat.—Among algae on rocky sub- strate. Perinereis villalobosi Rioja, 1947 Fig. 10 A-E Perinereis villalobosi Rioja, 1947: 532, figs. 18—22.—-Salazar Vallejo, 1989: 50. Material examined.—Baja California: Ensenada, Punta Banda, S. I. Salazar Val- lejo, coll., 7 Mar 1982 (1 specimen); Rin- con de Ballenas, V. Diaz Castafieda, coll., 20 Mar 1996 (4 specimens). Baja California Sur: La Paz, “El Comitan’’ beach, 1 km north of Centro de Investigaciones Biol6- gicas del Noroeste building, J. A. de Ledn- Gonzalez, coll., 1 Dec 1986 (12 speci- mens). Description. —Best preserved specimen 100 mm long and 4 mm wide including par- apodia, incomplete with 120 setigers. An- terior region with diffuse dark pigmentaton with no evident pattern. Prostomium as long as wide, with two pairs of eyes in trapezial arrangement, an- terior ones lensed; frontal antennae measur- ing half length of prostomium; palps glo- bose, with conical palpostyles. Peristomium as long as first two setigers; with longer PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tentacular cirri extending posteriorly to se- tiger 3 (Fig. 10A). Pharynx not everted in specimens. Anal- ysis performed by dissection. Paragnaths arranged on pharyngeal areas as follows: I, single cone; II, right side with 33 cones, left side with 24, both groups in triangular arrangement; III, 63 cones in 5 rows; IV, curved group of 74 cones; V, sin- gle large cone; VI, single short, transverse bar; VII—-VIII, 42 cones in 2 rows, but 1 row with 3 cones on each side. First two parapodia uniramous. Other anterior parapodia with rounded ligules; dorsal and ventral cirri well developed, dorsal ones better developed (Fig. 10B). On median and posterior parapodia noto- podial dorsal ligules enlarged and median ligules conical. Neuropodial postsetal lobes mamilliform, neuropodial ventral ligules capitate; dorsal cirri subdistally in- serted, ventral cirri proximally inserted (Bis “L0C, 'D).- All notosetae homogomph spinigers with slender, finely serrated appendage. Supra- cicular neurosetae homogomph spinigers and heterogomph falcigers (Fig. 10E). In- fracicular heterogomph spinigers and falci- gers. Discussion.—Perinereis villalobosi was described from Mazatlan, Sinaloa state, shores, from a single epitokous specimen. Hutchings et al. (1991), placed this species in their group “1A,” characterized by the presence of a transverse bar on Area VI and the notopodial dorsal ligule not greatly en- larged on middle and posterior segments. However, the transformation of posterior parapodia during the epitokous phase is such that it does not allow for the correct determination of the degree of enlargement of that ligule. The specimens reported in this study are referred to P. villalobosi due to their having the pharyngeal ornamenta- tion typical of the species. Since the noto- podial dorsal ligules are greatly enlarged, the species would belong to group “1B” of Hutchings et al. (1991). Distribution.—Eastern Pacific Ocean. Re- VOLUME 111, NUMBER 3 691 Fig. 10. Perinereis villalobosi. A. Anterior end, dorsal view; B. Parapodium 10th; C. Parapodium 49th; D. Parapodium 100th; E. Heterogomph neuropodial falciger in supracicular position from setiger 49. (Scale: A = 0.5 mm; B, C, D = 150um; E = 10pm). ported from the western coast of Baja Cali- fornia south to Mazatlan, Sinaloa, Mexico. Habitat.—In galleries in sandy rocks in supralittoral zone (found after fragmenta- tion of rocks). Key to the Species of Perinereis from Mexican Shores 1. Notopodial dorsal ligules enlarged — Notopodial dorsal ligules not enlarged. 692 Two cones in a row present on area I, short bar on Area VI ........ P. floridana 2. No bars on area IV — Two small basal bars on area IV in ad- dition to cones, short cone-shaped bar on ATE ANGIE wo Pe oe nen aus aces P. monterea 3. Single baronyareay Ville. ais eee 4 — Two bars onvarea VI ............... 7 4. sBawotearcayv short) sce eee ne 5 — Bar of area VI long and slender, ribbon- Shaped) eee, eS 5. ot TOR SRS 6 5. Areas I and V each with single cone. . LerA eR eR ee aM yirtda ees ees P. villalobosi — Area I with 4 cones; area V with 3 cones Sa ty tthe: anetapnedt a. ot ee Reet P. anderssoni 6. Area I with 7 cones; area VII—VIII with single row of 7 cones .. P. bajacalifornica — Area I with 11 cones; area VII-VIII with 37 cones in two rows P. elenaocasoae 7. Two cones on area I, no paragnaths on areas V; 11 cones in 2 rows on areas VT Vig he ee P. cariboea — Four cones in a diamond-shaped §ar- rangement on area I; single cone on area V; eight cones in single row on area VII— Vill P. osoriotafalli OO De O teoo Oo Ol Oo Oo ONO Acknowledgements We would like to thank K. Fauchald, and L. Ward (USNM) for their help in making available for examination the material un- der their care. Special thanks are also due to Leslie Harris, (LACM-AHFP), for kindly allowing us to examine the specimens from the AHF collections. Thanks are due to S. I. Salazar Vallejo who loaned us some ma- terial from the Mexican Caribbean area, to V. Diaz Castafieda who sent us the speci- mens of P. villalobosi from Rincon de Bal- lenas, Baja California; and to K. Sindemark (NRS), who sent us material of P. anders- soni. We also thank the two anonymous re- viewers whose comments greatly improved the manuscript. Literature Cited Augener, H. 1933. Polychaeten aus den Zoologischen Museen von Leiden und Amsterdam. II.—Zool- ogische Mededeelingen Uitgegeven Door’s PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rijks Museem van Natuurlijke Historie te Lei- den 16:107—128. Banse, K., & K. D. Hobson. 1974. Benthic errantiate polychaetes of British Columbia and Washing- ton.—Bulletin of Fisheries Research Board of Canada 185:1—111. Berkeley, E., & C. Berkeley. 1958. Some notes on a collection of Polychaeta from northeast Pacific south of Latitude 32°N.—Canadian Journal of Zoology 36:399—407. . 1960. Notes on some Polychaeta from the west coast of Mexico, Panama and Califor- nia.—Canadian Journal of Zoology 38:357— 362. Chamberlin, R. V. 1918. Polychaetes from Monterey Bay.—Proceedings of the Biological Society of Washington 31:173—180. Claparéde, E. 1870. Les Annélides Chétopodes du Golfe de Naples. Seconde partie —Mémoires de la Societé de Physique et d’Histoire Natu- relle de Genéve 20: 1—225, 31 pls, 365-542, 14 pls. Ehlers, E. 1868. Die Borstenwiirmer (Annelida: Chae- topoda) nach Systematischen und Anatomisch- en Untersuchungen Dargestellt. Leipzig. Wil- helm Engelmann. xxiv + 748 pp., 24 pls (pp. 1—268, pls. 1-11 published in 1864). Fauchald, K. 1977. Polychaetes from intertidal areas in Panama, with a review of previous shallow- water records.—Smithsonian Contributions to Zoology 221:1—76. Fauvel, P. 1930. Annélides Polychétes de Nouvelle Ca- lédonie recueillies par Mme A. Pruvot-Fol en 1928.—Archives de Zoologie Expérimentale et Générale 69:501—562. Grube, A. E. 1878. Annulata Semperiana. Beitrage zur Kenntniss der Anneliden Fauna der Philippinen nach den von Herrn Prof. Semper Mitgebrach- ten Sammlungen.—Mémoires de 1 Académie Impériale des Sciences de Saint Pétersbourg, ser. 7, 25(8):ix + 300, 15 pls. Hartman, O. 1951. The littoral marine annelids of the Gulf of Mexico.—Publications of the Institute of Marine Science 2(1):7—124. . 1968. Atlas of the Errantiate Polychaetous Annelids from California. Allan Hancock Foun- dation, University of Southern California, Los Angeles, 828 pp. Horst, R. 1889. Contributions towards the knowledge of the Annelida Polychaeta.—Notes from the Leyden Museum 11:38—45, 161-186, pls 3, 7, 8. . 1924. Polychaeta errantia of the Siboga Ex- pedition. Part III. Nereidae and Hesionidae.— Siboga Expedition Leyden 99 (Monograph 24): 145-198, 7 pls. Hutchings, P A., & A. Reid. 1990. The Nereididae from Australia. Gymnonereididae sensu Fitz- hugh, Ceratocephale, 1987: Australonereis, VOLUME 111, NUMBER 3 Dendronereides, Gymnonereis, Nicon, Olganer- eis, and Websterinereis.—Records of the Aus- tralian Museum 42(1):69—100. . , & R. S. Wilson. 1991. Perinereis (Polychaeta, Nereididae) from Australia, with redescriptions of six additional species.—Re- cords of the Australian Museum 43:241-—274. International Commission on Zoological Nomencla- ture. 1985. International Code of Zoological Nomenclature, Third Edition. International Trust for Zoological Nomenclature, London. 338 pp. Kinberg, J. G. H. 1866. Annulata nova.—Ofversigt af Forhandlingar Konglia Vetenskaps-Akadamiens 22:167-179,238-258. Kudenov, J. D. 1979. New species and records of Pol- ychaetous Annelids from the Tetraclita (Cirri- pedia: Crustacea) zone of the northern Gulf of California.—Bulletin of the Southern California Academy of Sciences 74(2):75-78. Linero-Arana, I. 1983. Dos nuevas especies de Nerei- dae (Polychaeta: Errantia) de la costa Oriental de Venezuela.—Boletin del Instituto Oceano- grafico de Venezuela, Universidad de Oriente 22(1 & 2):3-6. Monro, C. C. A. 1926. Polychaeta of the “Alert”? Ex- pedition. Families Hesionidae and Nereidae.— Zoological Journal of the Linnean Society 36: 311-323. Ramsay, L. N. G. 1914. Polychaeta of the family Ner- eidae, collected by the Scottish National Ant- arctic Expedition (1902—1904).—Transactions of the Royal Society of Edinburg 50:41—48, pl.3. 693 Rioja, E. 1947. Estudios Anelidolégicos XIX. Obser- vaciones sobre algunos nereidos de las costas de México.—Anales del Instituto de Biologia, México 18:527—535. . 1960. Estudios Anelidol6gicos XXIV. Adi- ciones a la fauna de anélidos poliquetos de las costas orientales de México.—Anales del Insti- tuto de Biologia, México, 31:289-316. Rozbaczylo, N., & J. C. Castilla. 1973. El género Per- inereis (Annelida, Polychaeta, Nereidae) en Chile.—Studies on the Neotropical Fauna 8: 215-232. Salazar Vallejo, S. I. 1985. Contribucién al conoci- miento de los poliquetos (Annelida: Polychaeta) de Bahia Concepcion, Baja California Sur. Tesis de Maestria, CICESE, 311 pp. . 1989. Enrique Rioja y su contribucion al es- tudio de los poliquetos (Annelida: Polychaeta) en México.—Brenesia 30:39—65. Salazar Vallejo, S. I., & M. S. Jiménez Cueto. 1996— 1997. Neréididos (Polychaeta) del Caribe Mex- icano con una clave para las especies del Gran Caribe.—Revista de Biologia Tropical 44(3)/ 45(1):361-377. Schmarda, L. 1861. Neue wirbellose Thiere beobachtet und gesammelt art einer Reise um die Erde 1853-1857. 1. Neue Turbellarien, Rotatorien und Anneliden.—Wilhelm Engelman, Leipzig 1(2):1-164, 22 pls. Treadwell, A. L., 1929. New species of polychaetous annelids in the collections of the American Mu- seum of Natural History, from Porto Rico, Flor- ida, Lower California, and British Somali- land.—American Museum Novitates 392:1—13. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):694—707. 1998. Streblospio gynobranchiata, a new spionid polychaete species (Annelida: Polychaeta) from Florida and the Gulf of Mexico with an analysis of phylogenetic relationships within the genus Streblospio Stanley A. Rice and Lisa A. Levin (SAR) Department of Biology, University of Tampa, Tampa, Florida 33606 U.S.A.; (LAL) Marine Life Research Group, Scripps Institution of Oceanography, La Jolla, California 92093-0218 U.S.A. Abstract.—Streblospio gynobranchiata, a new species of spionid polychaete, is described from Florida and the Gulf of Mexico. This species is distinguished by the presence of strap-like branchiae, beneath which larvae are brooded, in the genital body region of the females. Data from experimental crosses con- ducted in the laboratory are presented that indicate incipient reproductive iso- lation between S. gynobranchiata from Florida and S. benedicti from North Carolina and California. The spermatophores produced by males of S. gyno- branchiata and S. benedicti are described for the first time. Morphological, developmental, and reproductive characteristics are used in combination to con- struct a cladogram that suggests possible phylogenetic relationships between known species in the genus Streblospio. Genus Streblospio Webster, 1879 Diagnosis.—The genus Streblospio was established by Webster (1879) based upon material from New Jersey. Distinguishing features of the genus include: conical, rounded prostomium with 2—4 pairs of eyes and with or without a small nuchal tentacle; peristomium fused with the first setiger and forming a U-shaped envelope around the prostomium laterally and ventrally; one pair of palps and one pair of branchiae on the peristomium; a dorsal collar across the dor- sum of setiger 2; notosetae all capillary, neurosetae include capillaries, multidentate hooded hooks and ventral sabre setae; and pygidium simple, with ventral lappets or without appendages. Streblospio gynobranchiata, new species Figs. 1-13 Type material.—Holotype: adult female (USNM 178822), length 8.4 mm, Tampa Bay, Florida; Courtney Campbell Cause- way (west): depth 1 m, muddy sand, 27 Jul 1989. Paratypes: 17 males (USNM 178824), 8 females (USNM 178823), same collection data as holotype. Additional material prepared for scan- ning electron microscopy (SEM): Stub #1 (USNM 178825)—4 anterior ends, collect- ed 22 Jun 1987, Hillsborough River, Tam- pa, Florida, at the University of Tampa, depth 2 m, muddy sand; Stub #2 (USNM 178826)—2 adults, collected 23 Mar 1987, same as Stub #1; Stub #3 (USNM 178827)—4 anterior ends, collected 4 Aug 1988, same coll. data as holotype; Stub #5 (USNM_ 178828)—4 anterior ends, same coll. data as Stub #3; Stub #7 (USNM 178829)—4 adults, coll. 26 Mar 1987, same coll. data as holotype; Stub #8 (USNM 178830)—S5 adult females, coll. 2 May 1987, same coll. data as holotype; Stub #9 (USNM 178831)—S5 adults, same coll. data as Stub #8; Stub #10 (USNM 178832)—6 adults, same coll. data as Stub #8; Stub #11 (USNM 178833)—3 adults, coll. 1 Apr 1987, same coll. data as holo- type. VOLUME 111, NUMBER 3 yl / tif ; MWY a, V4 at’ § DY, 9), Figs. 1-4. Streblospio gynobranchiata, new species (USNM 178830): 1, whole female with characteristic brood structures; 2, posterior dorsal portion of female showing strap-like brood structures; 3, lateral view of brood structures in female; 4, dorsal view of most anterior brood structures and notosetae. Scales = 200 wm for 1; 100 pm for 2, 3. Other material examined.—Massachu- setts: Provincetown, coll. H. E. Webster, type (USNM 415), 6 specimens + 3 frag- ments; San Francisco Bay, Berkeley Beach, coll. O. Hartman, 27 Jul 1934, (USNM 20220) 8 paratypes; New Jersey, Little Egg Inlet, coll. 17 Apr 1972, (USNM 71677) 8 specimens, (USNM 71678) 22 specimens; SEM preparations: Box #7, Stub #4—1 an- terior end, Stub #6—3 anterior ends, coll. Bogue Sound, North Carolina, coll. L. Lev- in; Cultured material: Sebastian River, Flor- ida, 29 Mar 1993; Copano Bay, Texas, 18 Oct 1983; Godineaux, Trinidad 3 Apr 1984; Ballast Point, Tampa Bay, Florida 6 Jun 1984. Distribution.—Streblospio gynobran- chiata has been collected from the east and west coasts of Florida, Texas, and Trinidad. The type locality is Tampa Bay, Florida in the subtidal estuarine sediments of the Hillsborough River adjacent to the Univer- sity of Tampa in downtown Tampa. Etymology.—The specific name, gyno- branchiata, refers to the branchiate brood structures found in mature females of this species. The prefix is from the Greek, gyne, meaning woman or female plus the suffix from the Greek, branchia, refering to “gills.” Description.—Prostomium elongate and rounded anteriorly, flanked laterally by peristomial ridges leading to ventral mouth. One pair of palps located lateral to prostomium and median to peristomial ridges. Palps short; about same length as adjacent branchiae and densely ciliated on frontal surfaces (Figs. 1, 5). One pair of thick, ciliated branchiae inserted just pos- terior to palps. These branchiae with dis- 696 tal digitiform appendage terminally (Fig. 6). Small conical nuchal tentacle present between branchiae (Fig. 7). Nuchal ten- tacle difficult to see without removing branchiae. Notopodium of setiger 1 with 1-3 cap- illary setae and no postsetal lobe (Fig. 5). First neuropodium with 3—6 capillary setae and flap-like postsetal lobe. Second noto- podium with 6—12 capillary setae and broad, postsetal lobe continuous with dorsal hood that extends across dorsum connecting left and right notopodia. Anterior margin of hood flares upward and forms opening into dorsal pouch that extends posteriorly to end of third setiger (Fig. 5). Second neuropo- dium with 6-8 capillary setae and broad flap-like postsetal lobe. Notopodia of fe- males situated along lateral sides of body anterior to branchiate region but shifted dorsally throughout branchiate region (se- tigers 20—35) and on posterior setigers. No- tosetae in branchiate and posterior body regions longer than anterior notosetae. No- topodial lobes flap-like through anterior one-half of body, becoming digitiform and shifted dorsally toward posterior. Neurose- tae capillary in setigers 1—6. Ventralmost capillary seta in neuropodia 3—6 stouter than others and curving ventrally under body. Neuropodial hooded hooks and sabre setae beginning on setiger 7 and continuing to end of body (Figs. 8, 10). Hooded hooks accompanied by 1-3 small anterior capil- lary setae to near end of body. Single in- ferior sabre seta present in each neuropo- dium from setiger 7 to end of body (Fig. 8). Hooded hooks number 2-4 in setiger 7 to 6-8 near end of body; each hook with three or four pairs of small teeth in addition to main unpaired tooth (Fig. 10). Neuro- podial lobes flap-like on setigers 1—6, de- creasing in size posteriorly. Brood structures present on females from about setiger 20 to about 35 (24-35 in ho- lotype), small in anterior and posterior regions, reaching maximum size in middle of range (Figs. 1-4). Largest pouches com- posed of broad semicircular fold arising lat- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON erally between noto- and neuropodium and extending dorsally as thin digitiform ap- pendage resembling a branchium. Devel- oping larvae loosely held between branchi- ae by brooding females. Posterior to bran- chiate region on females are dorsal, seg- mentally arranged extensions of epidermis forming branched, ciliated appendages in rows between left and right notopodia (Fig. 9). Three or four of these extensions present on posterior setigers to near end of body. Cilia on these epidermal extensions ar- ranged in discrete tufts. Males of Streblospio gynobranchiata produce spermatophores in laboratory cul- tures in the presence or absence of females. These spermatophores are composed of a sickle-shaped proximal piece and a distal sperm ball (Fig. 11). The proximal piece is divided into two regions: a thick coarsely granular region (0.28 mm long and 0.12 mm wide) further from the sperm ball and a curved, conical, tapering region (0.24 mm long and 0.08 mm wide at widest point) leading to and attached to the sperm ball. The former, thick region has a groove down one side that ends at the junction between it and the conical region (Figs. 11, 13). The conical region is smoother than the thick region. The sperm ball measures 0.51 mm in diameter and is spherical in shape (Fig. 12). The dimensions of the spermatophore components vary with the size of the worm producing them, with larger worms releas- ing larger spermatophores. In cross-section, the material making up the thick and coni- cal regions of the spermatophore appears to be composed of fibrous material of un- known composition. The sperm ball is sur- rounded by a thin layer of this fibrous ma- terial. Only mature sperm are found inside the sperm ball. These sperm are oriented parallel to each other and closely packed inside the sperm ball with their tails ex- tending in a single direction. In fresh sper- matophores, the sperm tails have a refrin- gent color in reflected light and move in slow synchronous beating motions. Sper- matophores of the same general structure as VOLUME 111, NUMBER 3 697 Figs. 5-10. Streblospio gynobranchiata, new species (5, 9, USNM 178829; 6, USNM 178831; 7, USNM 178828; 8, 10, USNM 178830): 5, lateral view of anterior end with branchia (B), palp (P), and hood (arrow) visible; 6, distal tip of branchium showing finger-like extension (arrow): 7, lateral view of anterior end with right palp and branchium removed to expose the nuchal antenna (arrow); 8, lateral view of posterior setigers showing hooded hooks (H) and sabre setae (S), sabre setae are ventral to the hooded hooks; 9, dorsal view of female posterior to brood structures showing segmental epidermal extensions with tufts of cilia; 10, hooded hooks from posterior setiger showing pairs of small teeth distal to the main tooth. Scales = 100 wm for 5; 50 wm for 6, 7; 25 wm for 8, 9; 5 wm for 10. 698 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 11-13. Streblospio gynobranchiata, new species: 11, whole spermatophore as released by male; arrows indicate region enlarged in Fig. 13; 12, sperm ball attached to end of spermatophore; 13, central portion of spermatophore (region between arrows in Fig. 11) showing central groove and texture of material of spermato- phore. Scales = 100 pm for 11; 25 wm for 12, 13. described above were produced in labora- tory cultures of S. benedicti from North Carolina and Los Angeles. The mature sperm of S. gynobranchiata are greatly elongate with a mean head length (nucleus plus acrosome) of 41.15 wm (n = 150). Biology.—Streblospio gynobranchiata is found in estuarine to marine conditions in shallow-water sediment. Worms may form dense assemblages of tubes in stressed or disturbed habitats. Larval de- velopment is planktotrophic with brood- VOLUME 111, NUMBER 3 ing to the three-setiger stage followed by release into the plankton and settlement at about 9-12 setigers. Females that are forced to leave their tubes while brooding may drop their embryos prematurely. Male worms produce spermatophores (de- scribed above) that are incorporated into ventrally located seminal receptacles in the females. Site of fertilization and trans- port of mature oocytes into brood struc- tures have not been observed. Remarks.—In Webster’s (1879) original description of Streblospio benedicti, ref- erence is made to the genital pouches in females: “‘On one specimen the middle third of the dorsum was covered by a very thin, transparent, raised membrane.”’ (Webster 1879: 121). Webster (1879) did not use the term “‘pouches”’’ to describe these female structures; however, exami- nation of worms from Little Egg Inlet, New Jersey (USNM 71678) revealed brood pouches on females similar to those typically seen in worms from North Car- olina and California. Webster (1879) did not mention what have been called ‘‘sabre setae’’ in the neuropodium from setiger 7 to the end of the body nor did he describe the pairs of small teeth located above the main tooth on neuropodial hooded hooks. He refers to the hooded hook morphology as: “‘... they have four terminal teeth, the outer one being shortest ...’’ (Webster 1879: 121). Examination of the material from New Jersey (USNM 71678) by SEM revealed that sabre setae begin in setiger 7 and continue to near the end of the body and that the neuropodial hooded hooks have 3 or 4 pairs of small teeth in addition to the main unpaired tooth. The morpho- logical and reproductive characteristics of the species in the genus Streblospio are summarized in Table 1. Levin (1984) reported that Streblospio benedicti displayed different patterns of lar- val development over its range with some populations producing lecithotrophic larvae and others producing planktotrophic larvae. Planktotrophic and lecithotrophic popula- 699 tions were shown to be interfertile but fe- males from interpopulational crosses tended to produce larvae characteristic of their original population. The production of dif- ferent larval types was not affected by tem- perature or food availability in populations of S. benedicti from North Carolina (Levin & Creed 1986), although S. benedicti from Narragansett Bay increased brood sizes in response to elevated nutrient levels in me- socosm studies (Levin 1986). The demo- graphic consequences of planktotrophy ver- sus lecithotrophy in S. benedicti have been addressed by Levin, et al. (1987), Levin & Huggett (1990) and Levin & Bridges (1994). A genetic basis for this polymor- phism in S. benedicti was demonstrated by Levin et al. (1991). Buchanan (1890) described a species, Streblospio shrubsolii (originally Hekater- obranchus shrubsolii) from Sheppey, En- gland, that differed from S. benedicti by lacking a median nuchal tentacle (conical papilla or cirrus) on the first segment, lack- ing the conical dorsal cirri on posterior se- tigers, and lacking thoracic nephridia. Ad- ditional features of S. shrubsolii that distin- guish it from S. benedicti include: the first appearance of hooded hooks on setiger 8; the apparent lack of ventral sabre setae; the two-lobed dorsal hood on setiger 2; and the 2 rows of small teeth on hooded hooks. Subsequent examinations of this species re- vealed differences in egg diameter, location of the first gametogenic setiger, larval de- velopment, brood structures on females, and isozyme frequencies. According to Ca- zaux (1985) and our unpublished observa- tions (LAL), S. shrubsolii from France ex- hibit mature ova 200—230 pm in diameter, with one large larva brooded between trans- verse ridges on each segment. Development is direct in this species. In an examination of four enzyme systems (PHI, MDG-2, ADH, and IDH), individuals of Streblospio shrubsolii (n = 12) were fixed for com- pletely different isozymes than plankto- trophic individuals (n = 8-70) and leci- 700 Table 1.—Morphological and reproductive characteristics of described species of Streblospio. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Streblospio Streblospio Streblospio benedicti Streblospio benedicti shrubsolii japonica gynobranchiata Webster Buchanan Imajima Character new species 1879 1890 1990 Body length (mm) 8.4 6-12 6-10 7 No. of segments 70 70 48 48 Pairs of eyes 2-3 2-3 2-4 2 Nuchal antenna on Ist segment present present absent present Branchial pairs 1 1 1 1 Start of hooded hooks (setiger) 7 6/7/8/9 8 7 No. hooks per ramus anterior 3-4 3-5 2-3 5-6 posterior 8-10 8-12 5) u Dorsal collar one piece one piece 2 lobes one piece Brood structures branchiate pouches transverse, mid-seg- B mental ridges First saber seta 7 7 2 3 Rows of small teeth on hooded 3-4 34 De 5 pairs hooks Pygidium simple simple or lobed simple 2 lappets Egg diameter (wm) 70-90 70-90 (plankto- 200—230 ? trophic) 100—220 (lecitho- trophic) Larval development planktotrophic planktotrophic and _— Direct ? lecithotrophic Larval swimming setae yes yes (planktotrophic) no uy no (lecithotrophic) Number of larvae per brood 100—200 200—400 (plankto- ? ? trophic) 30-70 (lecitho- trophic) Sperm head length (wm) 41.15 47.81 ? y thotrophic individuals (n = 10—60) of S. the other species, as well as a different mor- benedicti. Studies of oogenesis in Streblospio have revealed differences in yolk bodies that ap- pear to vary with species and developmen- tal mode. The size of yolk granules and rel- ative proportion of heterosynthetically-de- rived yolk differs between planktotrophic and lecithotrophic forms of S. benedicti (Eckelbarger 1980, 1986). Preliminary ul- trastructural studies suggest that average yolk body diameter in S. shrubsolii (6.8 wm) is greater than that in S. gynobran- chiata (3.2 jm) from Fort Pierce, or from planktotrophic (2.6—-3.2 jm) or lecitho- trophic (3.2—5.3 ym) S. benedicti. The ma- ture egg of S. shrubsolii contains large quantities of lipid droplets not evident in phology of the egg envelope (K. Eckelbar- ger, pers. comm.). These differences sug- gest interspecific divergence in female ga- mete morphology similar to that reported for sibling species of Capitella (Eckelbar- ger & Grassle 1983). Horst (1909) described an additional spe- cies of Streblospio (S. dekhuyzeni) from Zuide Zee, France that had two types of capillary notosetae and more hooded hooks per ramus than S. shrubsolii. These differ- ences were minor enough to lead most tax- onomists to consider S. dekhuyzeni and S. shrubsolii synonyms (Fonseca-Genevois & Cazaux 1987). Hartman (1936) described Streblospio lu- tincola from several locations along the VOLUME 111, NUMBER 3 central California coast. She (pg. 46) con- sidered this new species to differ from S. benedicti in having a “... much smaller ventral peristomial fold, and its much larger prostomium.”’ Streblospio lutincola was l\at- er synonymized with S. benedicti (Hartman 1944). Foster (1971) proposed a synonymy of all previously described species making the genus Streblospio monotypic. In a note at the end of her treatment of the genus (pg. 115), she mentions a population of Streb- lospio collected from Maracaibo estuary that has branchiate brood structures in fe- males. This appears to be the first mention of S$. gynobranchiata in the literature. While Streblospio gynobranchiata gen- erally possesses a more southerly distri- bution than the other described species of Streblospio, it has been reported in the same geographical regions as S. benedicti. For example, lecithotrophic S. benedicti have been collected from Big Slough in Aransas Bay, Texas and S. gynobranchia- ta from an attached estuarine system, Co- pano Bay, Texas. Planktotrophic S. bene- dicti from Fort Pierce, Florida were col- lected within a few km of the Sebastian River system where S. gynobranchiata occurs. It is unknown whether these pat- terns of distribution are natural or the re- sult of anthropogenic transport since Streblospio may be transported in ship ballast water from one location to another (Carlton 1975). Imajima (1990) described a new subspe- cies (Streblospio benedicti japonica) from the Yatsu tidelands, Japan. This new form differs from S. benedicti in having ventral sabre setae beginning on setiger 3 and by having 5 pairs of small teeth in addition to the main unpaired tooth on the hooded hooks. The mean length of the sperm (acrosome + nucleus + middlepiece) from spermato- phores is significantly different (P < 0.001) between Streblospio gynobranchiata (41.15 wm + 4.22, n = 150) from Tampa Bay and S. benedicti (47.81 pm + 2.83, n = 150) 701 from Los Angeles. The location of the first gametogenic setiger in males of S. gyno- branchiata (15.66 + 9.98, n = 35) is not significantly different from that of plank- totrophs of S. benedicti from Fort Pierce (9.0 + 2.28, n = 6). In females of Streblospio gynobranchia- ta, the location of the first gametogenic se- tiger is significantly different from that of S. benedicti (planktotrophs and _ lecitho- trophs), and S. shrubsolii (Table 2). In ad- dition, the percent gametogenic setigers, the location of the first brood structure, and the total number of brood structures is signifi- cantly different between S. gynobranchiata and S. benedicti (Table 2). Reproductive isolation has been reported between Streblospio gynobranchiata from Tampa Bay, Florida and S. benedicti from North Carolina and from Los Angeles, Cal- ifornia (Rice 1991). Experimental crosses between populations were undertaken with worms that had been raised in isolation un- til sexual maturity then paired with a mate from an exotic population (experimental) or from the same population (control) and monitored for production of larvae. Fe- males from unsuccessful interpopulational crosses were subsequently paired with males from their own population to ensure female fertility. In 28 reciprocal crosses be- tween Tampa Bay and North Carolina worms, 26 crosses produced no offspring (see Rice 1991, table 11). In six crosses be- tween Tampa Bay females and Los Angeles males, no offspring were produced. In a separate series of experimental crosses, we were able to successfully cross Streblospio gynobranchiata from Copano Bay, Texas with lecithotrophic S. benedicti from Big Slough, Texas. We also found lar- val production in limited crosses between females of S. gynobranchiata from Tampa Bay and males (planktotrophic) of S. be- nedicti from Tar Landing, North Carolina, indicating that reproductive isolation be- tween these two (or more) species is not complete. In an experimental cross between one 702 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Reproductive characteristics of mature females of Streblospio species. Underlined mean values are not significantly different (P > 0.05). S. benedicti S. benedicti Analysis of Characteristic (planktotrophs) (lecithotrophs) S. gynobranchiata S. shrubsolii Variance Total setigers mean 47.8 45.0 46.6 51.3 F = 1.502 std. dev. 6.4 10.7 11.8 6.5 P = 0.2146 number 68 97 79 6 First gametogenic setiger mean 10.5 N27 Well 19 F = 217.2 std. dev. 1.1 1.3 2.0 1.1 P = 0.0001 number 71 80 78 6 % gametogenic setigers mean 38.3 34.6 45.4 — F = 12.7 std. dev. 8.7 6.8 8.7 — P = 0.0001 number 6 18 61 — First pouch/branchium mean 21.2 21.6 20.0 — F = 7.46 std. dev. Ded) 1.9 3.6 — P = 0.0007 number 69 94 66 — Total pouches/paired branchiae mean 9.6 8.4 14.5 — le = 16 .8)7/ std. dev. Del Dell MES) — P = 0.0001 number 69 92 66 —_ male of S. gynobranchiata from Copano Bay, Texas and one female of S. shrubsolii from Sully Island, Wales, larvae were pro- duced after 12 days but were misshapen and did not survive. In two experimental cross- es between males of S. gynobranchiata from Trinidad and planktotrophic females of S. benedicti from North Carolina, no lar- vae were produced. These results suggest that reproductive isolation is incipient be- tween S. gynobranchiata and S. benedicti and between S. gynobranchiata and S. shrubsolii. Reproductive compatibility be- tween populations is generally considered to be a pleisiomorphic characteristic, whereas loss of reproductive potential be- tween closely related taxa can be consid- ered an apomorphic characteristic. Consid- ering the degree of morphological differ- entiation (Tables 1, 2) in addition to the dif- ferences in geographic range and reproductive isolation, it seem certain that S. gynobranchiata represents a new and dis- tinct species. At present, there appear to be at least four species of Streblospio that can be dis- tinguished based upon morphological characteristics (Table 1). Streblospio gy- nobranchiata differs from S. benedicti in its more southern distribution, the pres- ence of branchiate brood structures on fe- males, the presence of posterior dorsal cil- iated appendages in females, gamete dis- tribution in females, and mature sperm di- mensions. Streblospio gynobranchiata differs from S. shrubsolii in the distribu- tion of hooded hooks (beginning on setig- er 7 in the former species and setiger 8 in the latter), the incomplete dorsal collar in the latter species, and the absence of a prostomial nuchal tentacle in the latter species. Streblospio gynobranchiata dif- fers from S. benedicti japonica in the dis- tribution of ventral sabre setae (beginning on setiger 3 in the latter species) and in the number of paired small teeth on the hooded hooks [see Ohwada & Nishino (1991) for a discussion of variability in VOLUME 111, NUMBER 3 this character]. In addition to the differ- ences in standard morphological charac- teristics between S. gynobranchiata and the other described species in the genus, there are also differences in gamete mor- phology, gamete distribution (Table 2), modes of larval development, and repro- ductive compatibility. Phylogenetic relationships.—The four species of Streblospio were analyzed for phylogenetic relationships using the cladis- tic programs, MacClade 3.01 (Maddison & Maddison 1992) and PAUP 3.1.1 (Swofford 1993). A character matrix consisting of 16 morphological and reproductive character- istics was constructed from literature re- ports and personal observations (Appendix 1). All characters were unweighted and unordered. The three most parsimonious trees found via an exhaustive search (Fig. 14A, B, C) all consist of 30 steps (consis- tency index = 0.933, retention index = 0.714) and were rooted using Paraprio- nospio pinnata and Spiophanes kroyeri as the designated outgroups. The strict consen- sus tree (Fig. 14D) groups S. gynobran- chiata and S. benedicti as a clade with S. benedicti japonica and S. shrubsolii joining as a polytomy. More information is needed concerning the characteristics of these latter two species in order to resolve this node. A bootstrap analysis of 100 replicates in PAUP using tree-bisection-reconnection branch swapping, MULPARS activated, zero-length branches collapsed, and multi- state characters interpreted as polymor- phism, resulted in the values listed on the branches in Fig. 14D. Sigvaldad6ttir et al. (1997) presented a cladistic analysis of the genera within the Spionidae based upon 25 morphological characters. Analysis of generic characteris- tics, including variation between species within a genus, resulted in a large number of most parsimonious trees (13305) and contributed little to our understanding of spionid relations. A second analysis using only type species as generic representatives produced more informative results with 703 four equally parsimonious trees. In the con- sensus tree derived from these four trees, Streblospio benedicti is grouped as the most plesiomorphic member of a clade including Aurospio dibranchiata, Orthoprionospio cir- riformia, Paraprionospio pinnata, Prion- ospio banyulensis, Prionospio steensrtupi, and Laubieriellus grasslei. Sigvaldad6ttir et al. (1997) point out that many characteris- tics of spionid genera are missing or incor- rectly reported in the literature and that res- olution of spionid phylogeny will likely re- quire re-examination of species and devel- opment of new characters. Incomplete literature reports and polymorphic charac- ters within species hampered resolution in the present study also. Four characters are coded as missing in the Streblospio data matrix for S. benedicti japonica since these four characters were not included in the de- scription of the species by Imajima (1990). Knowledge of these characters would likely help to resolve the polytomy in the Stre- blospio consensus tree. The study of variation within the genus Streblospio emphasizes the value of con- sidering reproductive characters along with standard morphological traits when distinguishing traits within a polychaete genus. Eckelbarger and Grassle (1984) demonstrated a similar situation in which sibling species of Capitella are distin- guished primarily by reproductive fea- tures. Characters such as strap-like bran- chiae in the genital region of females, the position and number of gametogenic se- tigers, and mature sperm dimensions dis- tinguish §. gynobranchiata from other species of Streblospio (Table 1). In mature specimens, reproductive characters like brood structures are more readily ob- served than setal or ciliation patterns. Un- fortunately, immature specimens of dif- ferent species of Streblospio appear sim- ilar in overt morphology and care must be taken with identification in regions where more than one species is likely to occur (e.g., southern USA, Europe). We expect that additional sibling species of Streblos- 704 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Streblospio gynobranchiata Streblospio benedicti Streblospio shrubsolii Streblospio benedicti japonica Spiophanes kroyeri Paraprionospio pinnata eect Streblospio gynobranchiata Streblospio benedicti Streblospio shrubsolii B Streblospio benedicti japonica Spiophanes kroyeri aioe: Paraprionospio pinnata Streblospio gynobranchiata Streblospio benedicti Streblospio benedicti japonica Streblospio shrubsolii Spiophanes kroyeri Paraprionospio pinnata 75 Streblospio gynobranchiata Streblospio benedicti 64 Streblospio shrubsolii Streblospio benedicti japonica Spiophanes kroyeri Paraprionospio pinnata Fig. 14. A, B, C, three most parsimonious trees for the four species in the genus Streblospio. Each tree is 30 steps; D, strict consensus tree with bootstrap values listed on branches. VOLUME 111, NUMBER 3 705, Table 3.—Character code data matrix. Taxon 1 2 3 4 5 6 7 8 Si Ok Pi Aah es 14 13 iG S. gynobranchiata 1 1 0) 1 0) 1 1 2 1 1 2 1 0) 0 1 1 S. benedicti 1 hy Or dO 1 1 2 I 0) 2 1 O 0/1 1 1 S. shrubsolii Oo 1 1 2 OV Oo ell 1 1 Dy ke OL eG 2 1 1 S. benedicti japonica 0 1 0) 1 0) v 1 2 1 20) ~2 1 ? 1 1 Paraprionospio pinnata 1 0) 1 3 1 DOO) SOLO) MRD = 3) OND 0) 1 1 Spiophanes kroyeri 0) 1 Q 4 2 1 OO 2a 2 1 R 2 y 0 O pio will emerge as reproductive, genetic, University of California Publications in Zoolo- and morphological characters of popula- By CIA 82. bits : : : . 1944. Polychaetous annelids from California tions around the world are examined in including the descriptions of two new genera greater detail. and nine new species.—Allan Hancock Pacific Expeditions. 10(2):239-307. Literature Cited Horst, R. 1909. De anneliden der Zuiderzee.—Mede- deelingen betreffende de uitkomsten der Zui- Buchanan, & 1890. Hekaterobranchus shrubsolii. A derzee—Expeditie, no. 5—Tijdschrift der Ned- new genus and species of the family Spioni- erlandsche Dierkundige Vereeniging, (series 2) dae.—Quarterly Journal of Microscopical Sci- II:138-152. ence 31:175—200. Imajima, M. 1990. Spionidae (Annelida, Polychaeta) Carlton, J. T. 1975. Introduced intertidal invertebrates. from Japan V. The genera Streblospio and Dis- Pp. 17—25 in R. I. Smith and J. T. Carlton, eds., pio.—Bulletin of the National Science Museum, Light’s Manual: Intertidal Invertebrates of the Series A (Zoology) 16:155—163. Central California Coast. University of Califor- Levin, L. A. 1984. Multiple patterns of development nia Press, Berkeley. in Streblospio benedicti Webster (Spionidae) Cazaux, C. 1985. Reproduction et développement lar- from three coasts of North America.—Biologi- vaire de l’annélide polychete saumatre Streb- cal Bulletin (Woods Hole) 166:498—508. lospio shrubsolii (Buchanan, 1890).—Cahiers . 1986. Effects of enrichment on reproduction de Biologie Marine 26:207—221. in the opportunistic polychaete Streblospio be- Eckelbarger, K. J. 1980. An ultrastructural study of nedicti (Webster): a mesocosm study.—Biolog- oogenesis in Streblospio benedicti (Spionidae), ical Bulletin (Woods Hole) 171:143—160. with remarks on diversity of vitellogenic mech- , & T. S. Bridges. 1994. Control and conse- anisms in Polychaeta——Zoomorphologie 94: quences of alternative developmental modes in 241-263. : a poecilogonous polychaete.—American Zool- . 1986. Vitellogenic mechanisms and the allo- ogist 34:323—332. cation of energy to offspring in polychaetes.— , & E. L. Creed. 1986. Effect of temperature Bulletin of Marine Science 39:426—443. and food availability on reproductive responses , & J. P Grassle. 1983. Ultrastructural differ- of Streblospio benedicti (Polychaeta: Spionidae) ences in the eggs and ovarian follicle cells of with planktotrophic or lecithotrophic develop- Capitella (Polychaeta) sibling species.—Bio- ment.—Marine Biology 92:103-113. logical Bulletin (Woods Hole) 165:379—393. , & D. V. Huggett. 1990. Implications of alter- , & . 1984. Value of life history studies native reproductive modes for seasonality and in the systematics of Capitella.—American Zo- demography in an estuarine polychaete.—Ecol- ologist 24:75A. ogy 71:2191—2208. Fonséca-Genevois, Ver6nica da, & C. Cazaux. 1987. , J. Zhu, & E. Creed. 1991. The genetic basis Streblospio benedicti Webster, 1879 (Annélide of life-history characters in a polychaete exhib- Polychéte) dans l’estuaire de la Loire: biologie iting planktotrophy and lecithotrophy.—Evolu- et écologie-—Cahiers de Biologie Marine 28: tion 45:380—397. 231-261. , H. Caswell, K. D. DePatra, & E. L. Creed. Foster, N. M. 1971. Spionidae (Polychaeta) of the Gulf 1987. Demographic consequences of larval de- of Mexico and the Caribbean Sea.—Studies on velopmental mode: Planktotrophy vs. lecitho- the Fauna of Cura¢gao and other Caribbean Is- trophy in Streblospio benedicti.mEcology lands 36(129):1-183. 68(6):1877-1886. Hartman, O. 1936. New species of Spionidae (Anne- Maddison, W. P., & D. R. Maddison. 1992. MacClade lida Polychaeta) from the coast of California.— 3: Analysis of phylogeny and character evolu- 706 tion. Sinauer Associates, Sunderland, Massa- chusetts. Ohwada, T., & Y. Nishino. 1991. Morphological vari- ation of hooded hooks in Prionospio japonica (Polychaeta, Spionidae).—Bulletin of Marine Science 48(2):246—250. Rice, S. A. 1991. Reproductive isolation in the Poly- dora ligni complex and the Streblospio bene- dicti complex (Polychaeta: Spionidae).—Bulle- tin of Marine Science 48:432—447. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sigvaldad6ttir, E., A. S. Mackie, & FE Pleijel. 1997. Generic interrelationships within the Spionidae (Annelida: Polychaeta),—Zoological Journal of the Linnean Society 119:473—500. Swofford, D. L. 1993. PAUP: Phylogenetic analysis using parsimony, version 3.1.1. Illinois National History Survey, Champaign, Illinois. Webster, H. E. 1879. Annelida Chaetopoda of New Jersey —Annual Report of the New York State Museum of Natural History 32:101—128, VOLUME 111, NUMBER 3 Appendix 1.—Character codes for phylogenetic analysis of Streblospio species (and Table 3). 16. 0 ils . Total number of segments <50 >70 . Membrane on palps present absent . Nuchal antenna present absent . First hooded hook 6th setiger 7th setiger 8th setiger Oth setiger 15th setiger . Pairs of Branchiae 1 pair 3 pair none Hooks per posterior ramus 5 8-12 >12 . Notopodial hooks present absent . Dorsal coilar at setiger 2 absent 2 lobes 1 lobe . Anterior postsetal lamella auricular semicircular pointed . Brood structures pouches branchiae none . First sabre seta 3rd setiger 4th setiger 7th setiger Oth setiger . Small teeth on hooded hooks 2 pair 3 or 4 pairs 5 pairs . Pygidium simple ventral lappets lobes and cirri Larval development planktotrophic lecithotrophic direct . Body regions distinct graded Neurosetal spines : present absent 707 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):708—719. 1998. Capitellids (Polychaeta: Capitellidae) from the continental shelf of the Gulf of California, México, with the description of a new species, Notomastus angelicae Pablo Hernandez-Alcantara and Vivianne Solis-Weiss Laboratorio de Ecologia Costera, Instituto de Ciencias del Mar y Limnologia, U.N.A.M., Apdo. Postal 70-305, Mexico, D.F 04510 Abstract.—In this study 11 species of capitellids in 6 genera are recorded from the continental shelf of the Gulf of California. The genus Notomastus is represented by 5 species, among them Notomastus angelicae is described. The Gulf of California is located approx- imately between 21°-32°N and 107° 114°W, in the northwestern region of Mex- ico. Shaped as a semiclosed basin, it has only one opening connecting it to the Pa- cific Ocean at its southern end. Due to its geographic location and the fact that several water masses are known to penetrate it (subarctic, tropical and subtropical) it is one of the most ecologically complex systems in Mexico. A large variety of habitats for the fauna (which can be locally highly di- versified) can be found there. The poly- chaetes and, in particular, the capitellid family is no exception to this: 28 species in 13 genera have been recorded so far in the continental shelf of the Gulf in addition to Six species from deep waters. Among these, 10 species in six genera were identified for this study, in addition to the new species, Notomastus angelicae, herein described. Capitellids are among the most common and widely distributed families of poly- chaetes worldwide. Found mainly in soft bottoms, they can occur in a large variety of sedimentary types and from intertidal to abyssal depths (Ewing 1984a). However, despite the relatively large number of spe- cies so far recorded in this family for the Gulf of California, our benthic studies of the region indicate that this is not an abun- dant family in the area (Hernandez-Alcan- tara 1992). Capitellids are known to be morpholog- ically relatively simple organisms which is partly why their correct taxonomic identi- fication is difficult. They were the first pol- ychaetes identified, Fabricius being the first to record in 1780 the species Capitella cap- itata (as Lumbricus capitatus). Since then, approximately 140 species in 36 genera have been described (Ewing 1984a). In the Gulf of California, the first record for this family was published by Fauvel (1943) who reported Dasybranchus caducus for San José Island (Baja California Sur). From the 34 species recorded so far, 12 have been originally described for the Gulf of Cali- fornia: Notomastus lobatus Hartman 1947, N. sonorae Kudenov 1975, N. angelicae n.sp., Dasybranchus parplatyceps Kudenov 1975, and D. platyceps Hartman 1947, in the continental shelf and Notomastus abys- salis Fauchald 1972, N. cinctus Fauchald 1972, Leiochrides hemipodus Hartman 1960, Neoheteromastus lineus Hartman 1960, Neomediomastus glabrus (Hartman 1960), Neonotomastus glabrus Fauchald 1972, and Notodasus magnus Fauchald 1972, in deep waters. This relatively high species richness can be partly explained both by considering the geographic situa- tion of the Gulf of California, which allows the confluence of tropical, subtropical and temperate waters, as well as its extended littorals with contrasting environmental conditions. VOLUME 111, NUMBER 3 Material and Methods The specimens were collected on board R/V El Puma (UNAM) as part of the insti- tutional ““CORTES”’ project in March 1985. A Smith-McIntyre grab was used to collect the sediment which was then sieved through a 0.5 mm sieve. Following fixation with 10% formalin, the organisms were separated, identified and preserved in 70% ethanol. The holotype of N. angelicae as well as some paratypes were deposited in the Na- tional Museum of Natural History, Smith- sonian Institution, Washington, D.C. (USNM) collection. Additional paratypes are deposited in Natural History Museum of Los Angeles County (LACM), Austra- lian Museum (AM), British Natural History Museum (BNHM), and in the Polychaete Collection of the Instituto de Ciencias del Mar y Limnologia, UNAM (CP-ICMyL), where the other identified species are also deposited. The habitat data for each species are ab- breviated as follows: D = depth in meters, T = temperature in °C, S = salinity in %o, OM = organic matter content in the sedi- ments in %, and DO = dissolved oxygen in m/l. Results In this study, 428 capitellids (11 species in six genera) were collected and identified from the continental shelf of the Gulf of California. Decamastus nudus Thomassin, 1970 Decamastus nudus Thomassin, 1970:81, figs. 6a—c. Material examined.—21 specimens: Ro- cas Consag, 30°59.4'N, 114°04.1’W, sta. 39, 106.4 m, silty sands, 16 Mar 1985 (1 spec- imen). North Tibur6n Island; 29°23.3’'N, 112°30.7'W, sta. 26, 71.9 m, fine sands, 14 Mar 1985 (7 specimens). San Miguel Cape, 28°08.0'N, 112°45.8’W, sta. 20, 54.1 m, fine sands, 13 Mar 1985 (10 specimens). Santa 709 Maria Bay; 25°02.4’N, 108°31.7’W, sta. 3, 32 m, fine sands, 10 Mar 1985 (3 speci- mens). Habitat.—Coarse substrates, coralline sands next to marine phanerogam meadows (Thomassin 1970). In this study it was col- lected in fine and silty sands; D = 32.0- 106.4; T = 13.2-14.4; S = 35.04—-35.35; OM = 1.5-5.7; DO = 1.02-3.25. Distribution.—Southwestern Madagascar (Thomassin 1970). North of Sonora, west of Santa Maria Bay, Sinaloa, and north of Santa Rosalia, Baja California Sur. Leiocapitella glabra Hartman, 1947 Leiocapitella glabra Hartman, 1947:438, pl. 54, figs. 1-3.—Ewing, 1984a:14.45, figs. 14.39, 14.40 a—d.—not Day, 1973: 100 type Ewing, 1984a. Material examined.—59 specimens: Ro- cas Consag, 30°59.4'N, 114°04.1’W, sta. 39, 106.4 m, silty sands 16 Mar 1985 (6 spec- imens). Tepoca Cape, 30°02.4’N, 112°55.4’W, sta. 44, 104.1 m, silty sands, 17 Mar 1985 (4 specimens). Punta Arboleda, 26°46.6'N, 110°06.7'W, sta. 14, 92 m, medium sands, 12 Mar 1985 (10 specimens). Santa Inés Bay, 26°59.4’N, 111°53.5’W, sta. 49B, 68.8 m, 19 Mar 1985 (26 specimens). Santa Inés Bay, 26°59.6'N, 111°50.4'W, sta. 49A, 100 m, fine sands, 19 Mar 1985 (13 specimens). Habitat.—53 to 100 m (Hartman 1947), silty clay sediments (Ewing 1984a). In this study it was collected in fine, medium and silty sands; D = 68.6—106.4; T = 13.2- 14.2; S = 35.09-35.26; OM = 3.0—7.2; DO = 0.92-2.40. Distribution.—Northern Gulf of Mexico (Ewing 1984a). Southern California; in the Gulf of California it has been reported in the Baja California eastern shores (Hartman 1947). Continental shelf of northern and southern Sonora, and west of Santa Inés Bay, Baja California Sur. Leiochrides hemipodus Hartman, 1960 Leiochrides hemipodus Hartman, 1960:136; 1963:22; 1969:381, fig. 1—\lFKauchald, 1972:242-—243. 710 Material examined.—1 specimen: Punta Arboleda, 26°46.6'N, 110°06.7’W, sta. 14, 92 m, medium sands, 12 Mar 1985 (1 spec- imen). Habitat.—Abyssal basins and canyons in muddy greenish sediments, with siliceous sponges (Hartman 1969). In this study it was collected in medium sands; D = 92.0; T = 13.6; S = 35.09; OM = 5.3; DO = 0.92. Distribution.—Southern California (Hartman 1969). In the Mexican Pacific, it has been collected in deep waters in front of Jalisco and Baja California (Hartman 1963), and Baja California Sur coasts (Fau- chald 1972). Punta Arboleda, Sonora, coasts. Mastobranchus ?variabilis Ewing, 1984 Mastobranchus sp. A.—Ewing, 1984a: 14.35, figs. 14.29, 14.30a-f. Mastobranchus variabilis Ewing, 1984b: 793-796, figs. la—d. Material examined.—6 specimens: El Fuerte River, 25°42.1’N, 109°30.6’W, sta. 51, 49.5 m, silty sands 20 Mar 1985 (2 spec- imens). Maria Madre Island, 21°38.2'N, 106°31.9’W, sta. 62C, fine sands, 22 Mar 1985 (4 specimens). Remarks.—The specimens analyzed are incomplete with a maximum of 41 setigers, which makes it impossible to confirm the presence of branchiae. According to the original description, they should be present in the middle-posterior region of the abdo- men as eversible “‘groups”’ of 5—8 cirriform filaments emerging from the posterior re- gion of the notopodial uncinigers. The pres- ence of capillary setae accompanying hood- ed hooks in most of the abdominal seg- ments also could not be ascertained. How- ever, the two other described species which belong to this genus, M. trinchesii Eisig 1887 and M. loii Gallardo 1968, have a bi- ramous first setiger, among other major dif- ferences with the specimens collected here. This led us to name the organisms as M. ?variabilis. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Habitat.—9.7 to 58 m, in sands (Ewing 1984b). In this study it was collected in fine and silty sands; D = 29.7—49.5; T = 14.8— 22.1; S = 35.10—35.15; OM = 4.2-—7.2; DO = 1.80—-5.29. Distribution.—Alabama and Florida, northern Gulf of Mexico, North Carolina shores (Ewing 1984b). West of El Fuerte River, Sinaloa, and Maria Madre Island, Nayarit. Mediomastus californiensis Hartman, 1944 Mediomastus californiensis Hartman, 1944: 264, pl. 6, figs. 64-65; 1947:408, pl. 46, figs. 3-4; 1969:387, figs. 1—4.—Day, 1973:99.—Hutchings & Rainer, 1979: 779.—Hobson & Banse, 1981:66.—Ew- ing, 1984a:14.14, figs. 14.9, 14.10a—c. Material examined.—2 specimens: Te- poca Cape, 30°08.6'N, 112°08.6'W, sta. 43, 68.8 m, silty sands 17 Mar 1985 (1 speci- men). Punta Willard, 30°11.5’N, 114°31.7'W, sta. 34, 32.9 m, sandy silty clay, 15 Mar 1985 (1 specimen). Habitat.—Intertidal to 517 m, commonly found in fine, muddy sands (Ewing 1984a). In this study it was collected in silty sands and muds; D = 32.9-68.8; T = 15.1—15.2; S = 35.38-35.45; OM = 6.9-8.9; DO = 3.03—4.30. Distribution.—Central and southern Cal- ifornia (Hartman 1969); from North Caro- lina to Florida; northern Gulf of Mexico, Australia (Ewing 1984a). Baja California coasts (Hartman 1963), Baja California Sur (De Leon-Gonzalez 1994), Sinaloa (Sala- zar-Vallejo 1981, Van Der Heiden & Hen- drickx 1982, Arias-Gonzalez 1984), and Jalisco (Varela-Hernandez 1993). Northern Gulf of California. Notomastus americanus Day, 1973 Notomastus americanus Day, 1973:100, figs. 131-n.—Ewing, 1984a:14.31, figs. 14.25,14.26a—d. Material examined.—61 specimens: Ro- cas Consag, 30°59.4’N, 114°04.1'W, sta. 39, VOLUME 111, NUMBER 3 106.4 m, silty sands, 16 Mar 1985 (12 spec- imens). Tepoca Cape, 30°02.4’N, 112°55.4'W, sta. 44, 104.1 m, silty sands, 17 Mar 1985 (8 specimens). Punta Willard, 30°11.5'N, 114°31.7'W, sta. 34, 32.9 m, sandy silty clay, 15 Mar 1985 (1 specimen). Punta Ar- boleda, 26°51.1’N, 110°06.5’W, sta. 15, 49.8 m, 12 Mar 1985 (13 specimens). Punta Arboleda, 26°46.6’N, 110°06.7’W, sta. 14, 92 m, medium sands, 12 Mar 1985 (1 spec- imen). El Fuerte River, 25°42.1'N, 109°30.6'W, sta. 51, 49.5 m, silty sands, 20 Mar 1985 (2 specimens). El Fuerte River, 25°46.8’N, 109°35.4’W, sta. 50, 97.0 m, sil- ty sands, 20 Mar 1985 (4 specimens). Santa Inés Bay, 26°59.6'N, 111°50.4'W, sta. 49A, 100 m, fine sands, 19 Mar 1985 (3 speci- mens). Punta San Marcial, 25°58.6’N, 111°06.9’W, sta. 10, 39 m, very fine sands, 11 Mar 1985 (4 specimens). Santa Maria Bay; 25°02.4'N, 108°31.7'W, sta. 3, 32 m, fine sands 10 Mar 1985 (1 specimen). Santa Maria Bay; 24°56.9'N, 108°41.8’W, sta. 4, 79 m, silty sands, 10 Mar 1985 (4 speci- mens). Maria Madre Island, 21°38.2'N, 106°31.9'W, sta. 62C, fine sands, 22 Mar 1985 (8 specimens). Remarks.—In small specimens, some- times we observed a mixture of hooks and acicular setae in the neuropodia of setiger 10, as Ewing (1984a) noted. In the same fashion, in some small organisms, the same feature can be observed but in the neuro- podia of eleventh setiger. Habitat.—35 to 100 m, in fine to very fine sands to coarse sands (Ewing 1984a). In this study, it was collected in fine, me- dium and silty sands, and muds; D = 29.7— 106.4; T = 13.2—22.1; S = 34.99-35.51; OM = 3.0-7.2; DO = 0.80-—5.29. Distribution.—North Carolina (Day 1973); northern Gulf of Mexico (Ewing 1984a). In the mexican Pacific it has been reported from the Jalisco coasts (Varela- Hernandez 1993), upper Gulf of California, southern Sonora, northern Sinaloa, and cen- tral Baja California Sur. 711 Notomastus hemipodus Hartman, 1945 Notomastus (Clistomastus) hemipodus Hartman, 1945:38; 1947:424, pl. 48, figs. 1-5; 1951:103, pl. 24, figs. 1-3; 1969: 393, figs. 1-5. Notomastus hemipodus.—Day, 1973: 100.—Ewing, 1984a:14.28, figs. 14.23, 14.24a—d. Material examined.—63 specimens: Ro- cas Consag, 31°16.1’N, 114°21.7'W, sta. 37, 30.3 m, fine sands, 16 Mar 1985 (11 specimens). Rocas Consag, 30°59.4’N, 114°04.1'W, sta. 39, 106.4 m, silty sands, 16 Mar 1985 (5 specimens). Punta Willard, 30°11.5'N, 114°31.7'W, sta. 34, 32.9 m, sandy silty clay, 15 Mar 1985 (2 speci- mens). Punta Arboleda, 26°53.2’N, 110°04.1’W, sta. 16, 22.2 m, fine sands, 12 Mar 1985 (3 specimens). Punta Arboleda, 26°51.1'N, 110°06.5'W, sta. 15, 49.8 m, 12 Mar 1985 (20 specimens). Santa Inés Bay, 26°59.6'N, 111°50.4’W, sta. 49A, 100 m, fine sands, 19 Mar 1985 (7 specimens). Punta San Marcial, 25°58.6’N, 111°06.9’W, sta. 10, 39 m, very fine sands, 11 Mar 1985 (13 specimens). Santa Maria Bay; 24°56.9'N, 108°41.8'W, sta. 4, 79 m, silty sands, 10 Mar 1985 (2 specimens). Habitat.—Intertidal to 120 m, in muds and muddy sands (Ewing 1984a). At 30 m and 30°C, 0.31% organic carbon, in sandy muds (Gonzalez-Ortiz 1994). In this study it was collected in fine, medium and silty sands and muds; D = 22.2—-106.4; T = 13.2-17.5; S = 35.00—35.51; OM = 2.4— 6.9; DO = 0.80—5.40. Distribution.—North Carolina (Day 1973); northern Gulf of Mexico (Ewing 1984a); southern California (Hartman 1969). In the mexican Pacific it has been collected in the Jalisco coasts (Varela-Her- nandez 1993), and the Gulf of Tehuantepec (Gonzalez-Ortiz 1994). Northern and cen- tral Gulf of California. Notomastus latericeus Sars, 1851 Notomastus latericeus.—Fauvel, 1927:143, figs. 49a—h.—Day, 1967:599, figs. 28.2a— 712 d.—Gallardo, 1968:120, pl. 53, fig. 13.— Thomassin, 1970:83, figs. 8a—-e.—Ewing, 1984a:14.26, figs. 14.29, 14.20a-e. Material examined.—20 specimens: Te- poca Cape, 30°02.4'N, 112°55.4’W, sta. 44, 104.1 m, silty sands, 17 Mar 1985 (5 specimens). Punta Willard, 30°11.5’N, 114°31.7'W, sta. 34, 32.4 m, sandy silty clay, 15 Mar 1985 (1 specimen). Estero Tastiota, 28°17.8’N, 111°37.1'W, sta. 47, 36.9 m, 18 Mar 1985 (1 specimen). Punta Arboleda, 26°51.1’N, 110°06.5’W, sta. 15, 49.8 m, 12 Mar 1985 (3 specimens). Santa Inés Bay, 26°59.4’N, 111°53.5’W, sta. 49B, 68.8 m, 19 Mar 1985 (1 specimen). Santa Maria Bay; 25°02.4’N, 108°31.7'W, sta. 3, 32 m, fine sands, 10 Mar 1985 (6 specimens). Banco Gorda, 23°06.6'N, 109°24.3W, sta. 56, 101 m, fine sands, 21 Mar 1985 (3 specimens). Habitat.—Intertidal to 4360 m; in very different kinds of sediments (Ewing 1984a). In this study it was collected in fine and silty sands and muds; D = 32.0—104.1; T = 13.7-15.1; S = 34.80—35.38; OM = 5.7— 7.2; DO = 1.02—4.30. Distribution.—Cosmopolitan (Ewing 1984a). Sonora coasts, northern Sinaloa, and northern and southern coasts of Baja California. Notomastus lineatus Claparede, 1870 Notomastus (Clistomastus) lineatus Clapa- réde, 1870:18, pl. 17, fig. 4—Hartman, 1947:419, pl. 46, figs. 1-2; 1969:395, figs. 1-5. Notomastus lineatus.—Fauvel, 1927:145, figs. 5la-1—Hobson and Banse, 1981: 66.—Ewing, 1984a:14.24, figs. 14.17, 14.18a-e. Material examined.—51 specimens: Te- poca Cape, 30°12.2'N, 112°46.9’W, sta. 42, 29.9 m, fine sands, 17 Mar 1985 (18 spec- imens). El Fuerte River, 25°39.9'N, 109°30.6’W, sta. 51, 49.5 m, silty sands, 20 Mar 1985 (1 specimen). San Miguel Cape, 28°10.4’N, 112°48.1’'W, sta. 19, 30.4 m, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON coarse sands, 13 Mar 1985 (14 specimens). San Miguel Cape, 28°08.0’N, 112°45.8'W, sta. 20, 54.1 m, fine sands, 13 Mar 1985 (3 specimens). Santa Inés Bay, 26°59.2’N, 111°58.3’W, sta. 49C, 28.9 m, very fine sands, 19 Mar 1985 (1 specimen). Punta San Marcial, 25°33.4’N, 110°59.8’W, sta. 8, 52 m, fine sands, 11 Mar 1985 (2 speci- mens). Punta San Marcial, 25°47.8'N, 111°03.8’W, sta. 9, 77.5 m, fine sands, 11 Mar 1985 (1 specimen). Banco Gorda, 23°08.7'N, 109°28.3’W, sta. 55, 32.5 m, fine sands, 21 Mar 1985 (11 specimens). Habitat.—Shallow subtidal to 298 m, in fine to coarse sands, sandy gravel and coral rubble (Ewing 1984a). In this study it was collected in fine, medium and coarse sands; D = 28.9-77.5; T = 13.6—21.3; S = 34.70— 35.54; OM = 1.8-7.2; DO = 1.80-—5.11. Distribution.—Mediterranean Sea (Fau- vel 1927); northern Gulf of Mexico (Ewing 1984a); Panama, Antarctic Ocean, from Canada to California (Hartman 1969). In the Mexican Pacific, reported from Todos los Santos Bay, Baja California Sur (Sala- zar-Vallejo 1985), and in Sinaloa coasts (Van Der Heiden & Hendrickx 1982). Mainly present in Baja California Sur lit- torals. Notomastus tenuis Moore, 1909 Notomastus (Clistomastus) tenuis.—Hart- man, 1947:420, pl. 47, figs. 1-5; 1969: 397, figs. 1-5. Notomastus tenuis.—Hobson & Banse, 198 1:66. Notomastus ? tenuis.—Ewing, 1984a:14— 26, figs. 14.21, 14.22a-e. Material examined.—100 specimens: Rocas Consag, 31°08.3’N, 114°13.3'W, sta. 38, 71.9 m, 16 Mar 1985 (4 specimens). Estero Tastiota, 28°16.4'N, 111°36.6'W, sta. 48, 60.2 m, fine sands, 18 Mar 1985 (5 specimens). Punta Arboleda, 26°46.6'N, 110°06.7'W, sta. 14, 92 m, medium sands, 12 Mar 1985 (5 specimens). El] Fuerte Riv- er, 25°39.9'N, 109°28.6'W, sta. 52, 28.6 m, silty sands, 20 Mar 1985 (5 specimens). El VOLUME 111, NUMBER 3 Fuerte River, 25°42.1’N, 109°30.6’W, sta. 51, 49.5 m, silty sands, 20 Mar 1985 (8 specimens). El Fuerte River, 25°46.8'N, 109°35.4'W, sta. 50, 97 m, silty sands, 20 Mar 1985 (5 specimens). San Miguel Cape, 28°07.7'N, 112°42.1'W, sta. 21, 104.1 m, 13 Mar 1985 (2 specimens). Santa Inés Bay, 26°59.2'N, 111°58.3'W, sta. 49C, 28.9 m, very fine sands, 19 Mar 1985 (5 speci- mens). Santa Maria Bay; 25°02.4'N, 108°31.7’W, sta. 3, 32 m, fine sands, 10 Mar 1985 (5 specimens). Santa Maria Bay; 24°54.6'N, 108°45.3’W, sta. 5, 120 m, silty sands, 10 Mar 1985 (2 specimens). Punta Mita, 20°53.9'N, 105°27.5'W, sta. 61, 50.4 m, fine sands, 23 Mar 1985 (6 specimens). Maria Madre Island, 21°38.2'N, 106°31.9'W, sta. 62C, 29.7 m, fine sands, 22 Mar 1985 (48 specimens). Habitat.—Intertidal to depths of 379 m, in muddy and sandy substrates (Hartman 1969). In this study it was collected in fine, medium and silty sands; D = 28.6—120.0; T = 12.9-22.1; S = 34.92-35.45; OM = 3.6-—7.2; DO = 0.54—5.40. Distribution.—From Canada to Califor- nia (Hartman 1969); northern Gulf of Mex- ico? (Ewing 1984a). In the Mexican Pacific, collected in the littorals of Baja California Sur (Rioja 1962, De Le6n-Gonzdélez 1994), Sinaloa (Salazar-Vallejo 1981, Van -Der Heiden & Hendrickx 1982), and in deeper zones from Baja California, Baja California Sur, Nayarit, and Jalisco (Fauchald 1972). It is found practically along the entire con- tinental shelf of the Gulf of California. Notomastus angelicae, new species Figs. la—d, 2, 3 Material examined.—44 specimens, Lo- cation: littoral, west of El] Fuerte River, Sin- aloa, México (25°39'54’N, 109°28'36"W), at a depth of 28.6 m. Holotype (USNM: 180697) and 5 paratypes (USNM: 180698); 5 paratypes (LACM-AHF-POLY-1902); 5 paratypes (A.M.: W24586); 5 paratypes (BNHM.: 1998.786-790); and 23 paratypes (CP-ICMyL: POP-17-001). 713 Description.—Holotype incomplete with 48 setigers, 15 mm long and 0.75 mm wide. Thorax length 3 mm. Paratypes with 21 to 40 setigers (mean = 29.48 + 5.43), 7.5 to 13 mm total length (mean = 10.0 + 2.12), and 0.75 to 1 mm width (mean = 0.76 + 0.12). Color of preserved specimens yellowish to light brown. Thorax slightly enlarged in first four to five setigers (Fig. 1a); epithelial surface clearly areolated as rhomboids in first four setigers (Fig. 2), following seti- gers smooth except for segmental biannu- lation. Inferior region of thorax from setig- ers 1-11 enlarged (Fig. 1b), as a conse- quence a middle longitudinal groove ap- pears to be present. Abdominal epithelium smooth. Prostomium short, a small triangular ter- minal palpode and two elongate ocular patches in posterior region (Fig. la). Peri- stomial ring achaetous, almost twice as long as following setigers. Eversible pharynx pa- pillose. Anterior thoracic notopodia dorsolateral, from setiger 7 widely separated. First seti- ger biramous. Only capillary setae in first ten setigers (10—20 per fascicle); last tho- racic segment (setiger 11) with capillary se- tae in notopodia and only hooded multiden- tate hooks (14—16) in neuropodia (Figs. Ic, 3))). Nephridial apertures located in segmental groove between each thoracic segment. Lat- eral organs present on all thoracic segments between noto- and neuropodium (Fig. 3). Transition from thorax to abdomen is ap- parent by size changes in segments, the presence of a hump-like protuberance in each segment and notopodial hooded hooks (Fig. la). Only multidentate hooded hooks in abdominal tori (Fig. 1d). Length of tori and hook number increase in neuropodia and disminish in notopodia towards poste- rior region. No branchiae. Abdomen with a large rounded dorso- transverse hump-like structure (Fig. le). Initially, tori located in middle part of seg- ments but migrating gradually towards pos- 714 C 0.02 mm 0.02 mm Bigeale PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ocular patches Notomastus angelicae, new species: a, thoracic and anterior abdminal region, lateral view; b, trans- verse section of fourth setiger; c, thoracic multidentate hooded hook; d, abdominal multidentate hooded hook; e, midabdominal segments, lateral view; f, pygidium of a regenerating organism. terior region of the segment until they al- most reach the intersegmental line. Pygidium unknown in all but one speci- men with 23 setigers well developed and a posterior regenerating region of 44 seg- ments with a pygidium. However, due to regeneration the shape could be somewhat atypical: it is tube-like with terminal part heart-shaped and a median digitiform cirrus (Fig. 1f). Habitat.—Specimens found at 28.6 m, in silty sands, 35.19 %o salinity, 16.8°C tem- VOLUME 111, NUMBER 3 Fig. 2. perature, 5.4 ml/l dissolved oxygen and 3.6% of organic matter. Distribution.—Notomastus angelicae is only present in the eastern coasts of the Gulf of California, west of El Fuerte River, Sinaloa. Etymology.—The species is named after Angélica Hernandez Huitron, niece of one of us (P H-A). Gender.—Feminine. Remarks.—Ewing (1984b) indicates that in the genus Notomastus, as well as in other capitellid genera, during setal development the thoracic hooks are gradually replaced by capillary setae in the middle-posterior region of thoracic neuropodia. As organisms reach adulthood, in some neuropodia a mixture of capillary setae and hooks appear; this arrangement is only SEM Prostomium and anterior thoracic region (peristomium) from Notomastus angelicae. present in one specimen analyzed with hooded hooks in setiger 10; the other 43 specimens only have hooks in neuropodia of setiger 11. Genus Notomastus Sars, 1851 was emended by Ewing (1982) to include cap- itellids with 11 thoracic setigers with only capillary setae on both rami or with neu- ropodial hooks in last 1—3 thoracic setigers. Notomastus angelicae differs from most species in the genus by having hooks in one thoracic neuropodium instead of only cap- illary setae in both thoracic rami. From the species with hooks in one or two thoracic neuropodia, none presents a prostomial pal- pode, and from those, N. precocis Hartman, 1960, N. teres Hartman, 1965, N. mossam- bicus (Thomassin 1970), N. americanus Day, 1973 and Notomastus sp. A Ewing, 716 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON / » mY > . ” i .) ok Bae! TG A! Sa 150 pu ge: 1984, all have only notopodia in first setiger (Table 1). Notomastus daueri Ewing, 1982, de- scribed from the northern Gulf of Mexico, is the only species in the genus with hooks in some thoracic neuropodia and first setig- er complete. Notomastus angelicae differs from it basically by the presence of the pro- stomial palpode, ocular spots, the abdomi- nal neuropodial shape and the well defined areolated area in the anterior thoracic re- SEM Setigers 10 and 11, lateral view from Notomastus angelicae. gion. The only species of the group with branchiae is N. daueri; however, these structures could not be seen here since the specimens are incomplete. Key to the Capitellids from the Gulf of California 1. Thorax with ten setigers — Thorax with more than ten setigers .. 3 2. Thoracic setigers with capillary setae VOLUME 111, NUMBER 3 10. only; first setiger incomplete, with no- tosetacron yam Decamastus nudus Setigers 1—4 with capillary setae; first setiger complete, with noto- and neu- MOSCUAG) Mediomastus californiensis mlonaxewithielleSetGens: aces oe 44 - 4 Thorax with more than 11 setigers .. 10 . Thoracic setigers with capillary setae only; abdominal notopodia with mixed fascicles of capillary setae and hooded hooks; first setiger incomplete ....... So bo Sn ee Mastobranchus ?variabilis Thoracic setigers with capillary setae only in both rami or with hooks only in neuropodia of last 1—3 setigers; abdom- inal setigers with hooded hooks only 20.0.8 i & MASc Sieh a) Hee ACE nen Mies Cee 5) Hirst setiser complete 2: (2257... 4. 6 First setiger incomplete ............ 8 Last thoracic segment with capillary se- tae only in both rami .............. Wf Last thoracic segment with capillary se- tae only in notopodia and hooded hooks Onlysineneuropodiay.. .- 5.0.6. -5..- oo Cea ee ae Notomastus angelicae Nephridial apertures restricted to tho- rax; thoracic segments uniannulate ... od 6 ae eae Notomastus lineatus Nephridial apertures restricted to ab- domen; thoracic segments biannulate 3 d,cto coke Set eae Notomastus latericeus All thoracic setigers with capillary se- RA CMOM aera: ned ety Pho tine Oo g) Last thoracic neuropodia with hooded hooks only ..... Notomastus americanus Prostomium with long ocular patches a Aker Pe Aedes, hie Notomastus tenuis Prostomium with pair of minute (often inconspicuous) eyespots ........... a bth cet ees eam ene ed Notomastus hemipodus Thorax with 12 setigers, all thoracic se- tigers with capillary setae only; first se- tiger incomplete .. Leiochrides hemipodus Thorax with 14 setigers; setigers 1-13 with capillary setae only in both rami, last thoracic setiger with capillary setae only in notopodia and hooded hooks only in neuropodia; first setiger incom- DIC es ria as, Leiocapitella glabra Acknowledgements We would like to thank especially Dr. Kristian Fauchald for his help in the revi- Table 1.—Morphological characters commonly used to differentiate the species of Notomatus with hooks in some thoracic notopodia. N. angelicae, new species Notomastus sp. A Ewing 1994 N. americanus Day 1973 N. mossambicus (Thomassin 1970) N. teres Hartman 1965 N. precocis Hartman 1960 N. daueri Ewing 1982 Character Present Absent Absent Absent Absent Absent Absent Prostomial palpode Present Present Present Present Absent Absent Absent Eyes or ocular spots First setiger Biramous Areolated Uniramous Plain Uniramous Plain Uniramous Areolated Uniramous Plain Uniramous Plain Biramous Anterior thoracic ep- Finely areolated ithelium Hooded hooks Branchiae Neuropodium 11 *Neuropodium 10-11 Absent *Neuropodium 11 Absent *Neuropodium 11 Absent Neuropodium 10-11 Absent Neuropodium 9-11 Absent Neuropodium 11 Present (around setiger 60) * In juveniles of these species hooded hooks or a mixture of capillary setae and hooks in setigers 9-11 can be present. WT 718 sion of the new species as well as his con- stant interest and help in our projects. Thanks are also due to M. Sc. Yolanda Homelas for the electron microscope pho- tographs. Literature Cited Arias-Gonzalez, E. 1984. Diversidad, distribuci6n y abundancia de anélidos (Poliquetos) en la Bahia de Mazatlan, Sinaloa, durante un ciclo anual. Tesis Profesional, Facultad de Ciencias, Univ- ersidad Nacional Autonoma de México, 102 pp. Claparede, E. 1870. Les Annélides Chétopodes du Golfe de Naples.—Mémoires de la Societé Phy- sique de Genéve 20:1—225. Day, J. H. 1967. A monograph on the polychaeta of Southem Africa. Part I: Errantia. Part II: Seden- taria.—British Museum of Natural History, Publication 656:1—878. . 1973. New Polychaeta from Beaufort, with a key to all species recorded from North Caroli- na.—wNational Marine Fisheries Service Circu- lar 375:1—140. De Le6n-Gonzalez, J. A. 1994. Poliquetos (Annelida: Polychaeta) de la plataforma continental de la costa oeste de Baja California Sur, México: tax- onomia, habitos alimenticios y distribucion. 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Fauna Groenlandica, systematice sistens, Animalia Groenlandica occidentalis hactenus indagata, quod nomen specificum, tri- viale, vernaculumque; synonyma auctorum plurium, descriptionem, locum, generationem, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mores, usum, capturamque singuli; pront dete- gendi occasio fuit, maximaque parti secundum proprias observationes. Hafniae et Lipsiae, 452 pp. Fauchald, K. 1972. Benthic polychaetous annelids from deep waters off western Mexico and ad- jacent areas in the eastern Pacific Ocean.—Al- lan Hancock Monographs in Marine Biology 7: 1-575. Fauvel, P. 1927. Polychétes sédentaires. Addenda aux Errantes, Archiannélides, Myzostomaires.— Faune de France 16:1—494. . 1943. Annélides polychétes de Californie re- cueillies par L. Diguet—Mémoires du Museum d’ Histoire Naturelle, Paris 18:1—32. Gallardo, V. 1968. Polychaeta from the Bay of Nha Trang, South Viet Nam.—NAGA Report, Sci- entific Results of Marine Investigations of the South China Sea and the Gulf of Thailand 1959-1961, 4(3):35-279. Gonzalez-Ortiz, L. 1994. Los poliquetos (Annelida: Polychaeta) de la plataforma continental del Golfo de Tehuantepec, México. Tesis Profesion- al, Facultad de Ciencias, Universidad Nacional Aut6noma de México, 191 pp. Hartman, O. 1944. Polychaetous annelids from Cali- fornia, including the description of two new genera and nine new species.—Allan Hancock Pacific Expeditions 10:239—307. . 1945. The marine annelids of North Caroli- na.—Duke University Marine Station, Bulletin 2:1—54. . 1947. Polychaetous annelids. Pt. 7. Capitelli- dae.—Allan Hancock Pacific Expeditions 10: 391-481. . 1951. The littoral marine annelids of the Gulf of Mexico.—Publications of the Institute of Marine Science 2:3—58. . 1960. Systematic account of some marine in- vertebrate animals from the deep basins off Southern California—Allan Hancock Pacific Expeditions 22:69—216. . 1963. Submarine canyons of Southern Cali- fornia. Part III. Systematics: Polychaetes.—AI- lan Hancock Pacific Expeditions 27(3):1—93. . 1965. Deep-water benthic polychaetous an- nelids off New England to Bermuda and other North Atlantic areas ——Allan Hancock Founda- tion Publications Occasional Papers 28:1—378. . 1969. Atlas of the Sedentariate Polychaetous Annelids from California. Allan Hancock Foun- dation, University of Southern California, Los Angeles, 812 pp. Hernandez-Alcantara, P. 1992. Los poliquetos (Anne- lida: Polychaeta) de la plataforma continental del Golfo de California, México. Taxonomia, abundancia numérica y distribucién geografica. VOLUME 111, NUMBER 3 Tesis Maestria, UACPyP-CCH, Universidad Nacional Aut6noma de México, 427 pp. Hobson, K. D., & K. Banse. 1981. Sedentariate and archiannelid polychaetes of British Columbia and Washington.—Canadian Bulletin of Fish- eries and Aquatic Sciences 209:1—144. Hutchings, P. A., & S. FE Rainer. 1979. The polychaete fauna of Careel Bay, Pittwater, New South Wales, Australia—Journal of Natural History 13:745-796. Kudenov, J. D. 1975. Sedentary polychaetes from the Gulf of California, Mexico.—Journal of Natural History 9:205—231. Moore, J. P. 1909. Polychaetous annelids from Mon- terey Bay and San Diego, California.—Proceed- ings of the Academy of Natural Sciences of Philadelphia 61:235—295. Rioja, E. 1962. Estudios anelidol6gicos XX VI. Algu- nos anélidos poliquetos de las costas del Paci- fico de México.—Anales del Instituto de Biol- ogia, Universidad Nacional Aut6noma de Méx- ico 33:131—229. Salazar-Vallejo, S. I. 1981. La coleccié6n de poliquetos (Annelida: Polychaeta) de la Facultad de Cien- cias Bioldgicas de la Universidad Auténoma de 719 Nuevo Leon, México. Tesis Profesional, Facul- tad de Ciencias Bioldgicas, Universidad Autén- oma de Nuevo Leon, 156 pp. . 1985. Contribucién al conocimiento de los poliquetos (Annelida: Polychaeta) de Bahia Concepcion, Baja California Sur, México. Tesis Maestria, Centro de Investigaciones y Estudios Superiores de Ensenada, México, 311 pp. Sars, M. 1851. Beretning om en y Sommeren 1849 Foretagen Zoologisk Reise y Lofoten og Fin- marken.—Nyt Mag. Naturvid. Oslo 6:121—211. Thomassin, B. 1970. Contribution a |’étude des poly- chétes de la région de Tuléar (SW de Madagas- car). Sur les Capitellidae des sables coral- liens.—Recueil des Travaux de la Station Ma- rine d’Endoume, Supplément 10:71—101. Van Der Heiden, A. M., & M. E. Hendrickx. 1982. Inventario de la fauna marina y costera del sur de Sinaloa, México. Segundo Informe, Instituto de Ciencias del Mar y Limologia, Universidad Nacional Aut6noma de México, 135 pp. Varela-Hernandez, J. J. 1993. Anélidos poliquetos de la plataforma continental de Jalisco, México. Tesis Profesional, Facultad de Ciencias Biol6- gicas, Universidad de Guadalajara, 113 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):720—730. 1998. Phytoplankton composition within the tidal freshwater region of the James River, Virginia Harold G. Marshall and Lubomira Burchardt (HGM) Department of Biological Sciences, Old Dominion University, Norfolk, Virginia 23529-0266, U.S.A.; (LB) Department of Hydrobiology, Adam Mickiewicz University, Marcelinska 4, 60-801, Poznan, Poland Abstract.—Based on a 10.5 year data set, 271 taxa were identified at a single tidal freshwater station in the James River. The mean monthly concentrations of major algal categories, total biomass and productivity are given. Diatom maxima were associated with peak periods of river discharge, with chloro- phytes, cyanobacteria, and autotrophic picoplankton abundance and productiv- ity greater during reduced river flow and more stable water conditions. Within the tidal stretch of estuarine riv- ers, there is a unique and little studied re- gion dominated by freshwater plankton, yet it is subjected to daily tidal influence and the occasional intrusion of estuarine spe- cies. These waters are referred to as tidal fresh. The channel divisions between fresh- water, tidal fresh, and oligohaline sections of a river are not constant, but will move longitudinally within the river basin in re- sponse to changes in the amount of river flow and tidal influence. During a period of drought, the range of the tidal fresh area moves farther upstream, whereas, during the seasonal rains, it would extend farther downstream. The abundance of tidal fresh algae in this region has been associated to hydrodynamic events in the river, with cell concentrations inversely related to fresh- water input and directly related to the wa- ter’s residency time (Jackson et al. 1987, Schuchardt & Schirmer 1991, Jones et al. 1992). Other major environmental factors influencing abundance in this region in- clude turbidity, nutrients, tidal cycles, and light availability (Cole et al. 1982, Cloern et al. 1983, Cloern 1987, Haas et al. 1981, Jones 1991). The extent of dominance by freshwater algae downstream varies within different estuaries, but the downstream flora will be dominated by estuarine species (Haerte et al. 1969, Forester 1973, Jackson et al. 1987, Marshall & Alden 1990). In tid- al freshwater (<0.5 ppt), the algae are mainly composed of chlorophytes, cyano- bacteria, and diatoms (Forester 1973, Opute 1990, Marshall 1994), with diatoms often predominating (Haertel et al. 1969, Jackson et al. 1987, Schuchardt & Schirmer 1991). Diatoms are more common during periods of high river discharge in contrast to chlo- rophytes, cyanobacteria, and phytoflagella- tes, which are more dominant during low river discharge (Farrell 1994, Schmidt 1994). The James River is a major tributary of the Chesapeake Bay, with a drainage basin of 26,440 km’, a length of 545 km and a mean monthly discharge rate of approxi- mately 7100 ft? sec”! (Belval et al. 1995). Within the southern Chesapeake Bay trib- utaries, several studies have compared phy- toplankton spatial and temporal distribution within the tidal fresh and saline sections of these rivers (Marshall & Alden 1990, Mar- shall & Affronti 1992, Marshall 1994). These studies indicate the transport of es- tuarine species upstream via sub-surface waters and the transition to estuarine spe- cies dominance downstream. Marshall and VOLUME 111, NUMBER 3 Alden (1990) recognized two distinct and dominant assemblages within the tidal James River. These were tidal fresh and oli- go-mesohaline populations. The tidal fresh- water group was dominated by the centric diatoms Skeletonema potamos, Cyclotella striata, and several Aulacoseira spp. Estu- arine diatoms became dominant down- stream. These included Skeletonema costa- tum, Leptocylindrus minimus, and Cyclotel- la caspia (C. choctawhatcheeana). In the James River, Filardo and Dunstan (1985) reported productivity in the lower saline regions was inversely correlated to river discharge. They noted inverse relationships between phytoplankton abundance in the oligohaline reach of the river to phyto- plankton biomass downstream, suggesting this region may control nutrient availability downstream. Other studies within Chesa- peake Bay tributaries have indicated phy- toplankton development may become nutri- ent limited in spring due to reduced phos- phorus and silicon levels, and in late sum- mer as nitrogen levels decrease (Anderson 1986, Webb 1988). Based on a 5-year study of the James River, Belval et al. (1995) re- ported median annual concentrations near the fall line of total nitrogen (TN) and total phosphorus (TP) as 0.74 and 0.15 mg 1"', respectively. The median value for dis- solved orthophosphate was 0.04 mg 1". They found total nitrogen, total phosphorus, and total suspended solids (TSS) directly related to increase river discharge, and dis- solved orthophosphate increased with peri- ods of reduced river flow (and summer). Mean annual productivity for the tidal freshwater station (TF5.5) in the James Riv- er was reported by Marshall & Nesius (1993) as 279.9 gC m* yr'', with higher values noted downstream. Using a 5-year data set for the station in this study (TF5.5), they gave mean water concentrations for the following parameters (in mg 17!: Si (5.8), O, (8.5), TP (0.16), TN (0.89), and TSS (30.3), and a mean pH of 7.5. In the tidal freshwater section of the Potomac Riv- er, Jones (1991) associated high photosyn- 721 thesis rates during cyanobacteria blooms to increased pH (>9) and the release of sedi- ment phosphorus into the water column, which may then stimulate more productiv- ity. The objectives of this study are to: 1) identify phytoplankton taxa at a tidal fresh- water station in the James River, and 2) present the mean monthly concentration levels for the dominant phytoplankton cat- egories in this region. This information will come from a 10.5-year data base at this sta- tion. Methods Phytoplankton composition and abun- dance were determined from a vertical se- ries from surface to bottom of monthly composite water samples taken from July 1986 through December 1996 at Station TF5.5 in the James River as part of the Chesapeake Bay Plankton Monitoring Pro- gram (Marshall & Alden 1990). The station is located above the turbidity maximum zone (37°18'46"N, 77°13'59"W), with mean water depth of 10.6 m, and a tidal range of <1 m (Marshall 1994). Monthly 500 ml sub-samples were taken for analysis using a modified Utermohl method to settle, si- phon, and concentrate the water sample into settling chambers for microscope analysis (Marshall & Alden 1990). Identification and cell abundance for each sample were based on a minimum microscope count of 200 cells, using a minimum of 10 random fields examined at both 315X and 500%, in addition to having the entire bottom of the settling chamber scanned at 125% for larg- er, less abundant cells that may be missed in the random field counts. The autotrophic picoplankton component and productivity values are based on a 7.5 year data set de- rived from monthly samples taken from July 1989 through December 1996. The autotrophic picoplankton were distin- guished from heterotrophic cells using epi- fluorescence microscopy (Marshall 1995), with C productivity measurements taken 122; £ © oo © Thousand Cubic Feet/Second = ND ro) oO J F M A M High-low and mean monthly river discharge rates in the James River (fall line values), 1986 through Fig. 1. 1996. as described by Marshall and Nesius (1993). A 12-month set of samples from 1995 were also re-examined for further spe- cies identification using both light and scan- ning electron microscopy. Biomass was cal- culated from cell volume measurements as cell carbon (Smayda 1978). Reference to seasons uses December, January and Feb- ruary as winter, followed by the sequential 3-month periods for spring, summer, and fall. Water discharge rates were provided through the information data bank of the U.S. Geological Survey. Results The tidal fresh station (TF5.5) main- tained <0.5 ppt salinity during this study. A total of 271 taxa were identified within the following categories: Bacillariophyceae (78), Chlorophyceae (94), cyanobacteria (36), Xanthophyceae (19), Euglenophyceae (15), Dinophyceae (13), Chrysophyceae (9), and Cryptophyceae (7). The autotro- phic picoplankton were collectively com- James River Discharge 1986-1996 High-Low Range with Mean Flow PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON J A -S “Oo “Nees piled as a composite group, but consisted mainly of single-celled cyanobacteria, but these are not included in the cyanobacteria concentrations given in Fig. 4. The pattern of river discharge in the James river is maximum flow in spring, re- duced discharge during summer, and an in- crease during fall and winter (Fig. 1). The greatest monthly ranges for flow occurred during the period of spring rains in March and April, and least during summer. During the 10.5-year study, annual mean monthly flows ranged from 4495 (1988) to 13,192 (1996) ft? sec~'. Many of the high monthly ranges represented extreme, and not com- mon occurrences; therefore the mean flow rate is considered here more typical for each month. The most consistent and least variable months of flow were July and Au- gust. During these months the salinity gra- dients would move farther upstream and more stratified water column conditions would exist in the river, in contrast to the high discharge periods of spring. Consid- VOLUME 111, NUMBER 3 Bae8 || (Aae See il | -- =e EEEEEET. MA M Fig. 2. December 1996. erable variability in the amount and timing of river discharge occurred, which influ- enced the residency time and period of de- velopment for phytoplankton in the water column. The mean monthly productivity for this site is given in Fig. 2. The period of highest productivity occurred between mid-spring (April) and continued into mid-fall (Octo- ber). These values decreased into winter. The lowest production levels were in Jan- uary at 2.3 mgC m? hr“! (Fig. 2). In sum- mer, they reached 71—77 mgC m? hr !. This period coincided with maximum develop- ment of the total phytoplankton and auto- trophic picoplankton components. The spring and fall pulses were dominated by diatoms, whereas the summer months con- tained a diverse assemblage of phytoplank- ton that contributed to this productivity. The phytoplankton composition was dominated by freshwater diatoms, chloro- phytes, cyanobacteria, and cryptomonads. Maximum total phytoplankton abundance occurred during April (64 X 10° cells 1~') and from July through October (79-109 x Ute GAS eS Mean monthly productivity rates at the tidal freshwater station (TF5.5) from July 1989 through 10° cells 1~'), after which came a sharp de- crease into winter (Figs. 3, 4). The spring development was a product of the diatom pulse, whereas a combination of chloro- phytes, diatoms, and cyanobacteria were mainly responsible for the summer and fall growth. The mean monthly biomass pattern was similar to phytoplankton concentra- tions, having greater biomass occurring in spring and from mid-summer through fall (Fig. 3). In addition to the primary cate- gories responsible for the cellular abun- dance mentioned above, the biomass levels were enhanced by euglenophytes during their peak time of development in July and August (Fig. 4). Bacillariophyceae.—The lowest diatom concentrations (3 X 10° cells 1~') were in winter and early spring (Fig. 3). Cell num- bers increased with peak abundance occur- ring during the spring pulse (April and May), when mean concentrations reached 41 X 10° cells 1-'. The maximum diatom development also coincided with the peak discharge period of April. There were re- duced diatom concentrations in summer, 724 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Millions Millions 120 Cells/liter J F M A M J J A S O N OD Millions Millions 7 ian) SL fame a BE eh neal (SE Lan] oe Seer ea Pl tl Pe Fig. 3. Mean monthly abundance for total phytoplankton, diatoms, and chlorophytes, and the mean monthly phytoplankton biomass at the tidal freshwater station, (TF5.5), July 1986—December 1996. Millions -[ervetepnyesse [Ta Millions “oer TTT Cells/liter =" Euglenophyceae ’®i{Autotrophie Picoplankton |/ | |_| J RAM, AS Moede cdi vA, S)-.0))\N.—D Fig. 4. Mean monthly abundance at the tidal freshwater station (TF5.5) for cyanobacteria, cryptophytes, and euglenoids from July 1986—December 1996, and for the autotrophic picoplankton July 1989—December 1996. VOLUME 111, NUMBER 3 during a time of reduced river flow, which was then followed by a less developed fall pulse (20-25 X 10° cells 1~') in October. The species most responsible for the spring and fall maxima were freshwater centric di- atoms: Actinocyclus normanii, Aulacoseira granulata, A. granulata v. angustissima, Cyclotella striata, C. meneghiniana, Cy- clostephanos spp., Melosira varians, Ske- letonema potamos, Stephanodiscus hantz- schii, and Thalassiosira lacustris. Of these, Skeletonema potamos was most abundant during seasonal maxima and present throughout the year. In addition, the abun- dant pennate diatoms included Cymbella af- finis, Nitzschia acicularis, several other Nitzschia spp., Staurosirella leptostauron, and Surirella elegans. Many of these cen- tric and pennate species have also been re- corded downstream in the James River, but decreasing in their abundance with in- creased salinity (Marshall 1994). The ma- jority of the estuarine diatoms recorded for this site were noted in <2% of the samples. More frequently recorded (13-19% of the collections) were Skeletonema costatum, Leptocylindrus danicus, and L. minimus. Chlorophyceae.—The chlorophytes had the largest number of taxa (94) among the algae, with their greatest abundance in sum- mer and fall (Fig. 3). Development coincid- ed with the decrease of the spring diatom bloom, reduced river flow, and accompa- nied the increase of cyanobacteria. Highest numbers were recorded during this period (4-5 X 10° cells 1~'), with reduced concen- trations during winter and spring. The most widely represented genera included: Ankis- trodesmus, Crucigenia, Kirchneriella, Mon- oraphidium, Scenedesmus, Staurastrum, Tetraedron, and Tetrastrum. None of the chlorophytes dominated the phytoplankton in abundance; however, they were common constituents of the algal community throughout the year. Cyanobacteria.—This category was rep- resented by a variety of filamentous and co- lonial species that were most abundant (Fig. 4) from early summer (June) to mid-fall 725 (October). Their peak development was at 50 X 10° cells 1~' (August—October). Cell abundance then declined rapidly into winter and remained low through spring. The most common species throughout the year were Chroococcus minor, Dactylococcopsis aci- cularis, D. raphidioides, Merismopedia punctata, Oscillatoria agardhii, and O. lim- netica. These taxa were major contributors to the summer—fall maximum. Other spe- cies in abundance included Anabaena soli- taria, A. affinis, Gomphosphaeria lacustris, Merismopedia tenuissima, and M. glauca. This group was associated with a more stratified water column, increased water temperatures, and reduced river discharge. Cryptophyceae.—The cryptomonads represented a ubiquitous and abundant component of the phytoplankton assem- blages throughout the year, and are also common in the downstream tidal sectors of the James River (Marshall & Alden 1990). Their mean monthly concentrations for the year ranged from 5—15 X 10° cells 1~' (Fig. 4). They were most abundant in winter (Dec.), late spring (May), and mid-summer (July). Cryptomonas marssoni was present during each season. Other widely distrib- uted species were Cryptomonas ovata and Rhodomonas minuta. The peak abundance of cryptomonads was associated with re- duced river discharge and summer. Autotrophic picoplankton.—The autotro- phic picoplankton consisted of mainly sin- gle cell cyanobacteria 0.5—1.5 ym in size (e.g. Synechococcus sp.) and represent a ubiquitous component of the James River and Chesapeake Bay (Marshall & Nesius 1993, Marshall 1995). Their peak abun- dance was during the summer months at 26-60 X 10’ cells 1~' (Fig. 4). Their lowest concentrations occurred in February and March (3-5 X 10° cells 1~'). Some chlo- rophytes were also present in this group, but they represented only a small fraction of the composition. A major importance of the autotrophic picoplankton in the James River and the lower Chesapeake Bay is 726 their contribution to the total summer pro- ductivity (Marshall & Nesius 1993). Other categories.—The euglenophy- ceans were generally found in low concen- trations throughout the year, with the ex- ception of July and August, when their mean concentrations reached 11 X 10° cells 1~' (Fig. 4). Common taxa included Eugle- na viridis and E. oxyuris, with other Eugle- na, Phacus, Trachelomonas, and Strom- bomonas species less abundant. The xan- thophytes were recorded throughout the year. However, they were generally noted in low concentrations, with low monthly spe- cies diversity. Tribonema monochloris was the most common species. Several species were associated with increased summer and fall development. These included Dicho- romococcus curvatus and Pseudotetraedron neglectum, whereas Botrydiopsis eriensis and B. arhiza were noted in spring and summer. Other species within this category were less common. The chrysophytes were represented by 5 Dinobryon species, Synura uvella, Lagy- nion cystodini, and Dictyota fibula. Dino- bryon sociale was most frequently noted in late winter and early spring. Dictyota fibula was rare (1% of the samples), coming from an infrequent intrusion of downstream wa- ter into this region. The dinoflagellate pop- ulations were also low, with their counts en- hanced by occasional sporadic growth pe- riods. The maximum development of di- noflagellates occurred in winter (January) and between late spring (June) and fall (Au- gust), with mean concentrations of 2-6 X 10* cells 1~'! during summer months. There was low species diversity of dinoflagellates throughout the year, with a limited number of dominant taxa. The genera most fre- quently identified were Gymnodinium, Gy- rodinium, Peridinium, and Prorocentrum. Downstream species that occurred within these collections included Heterocapsa tri- quetra, Prorocentrum micans, and P. min- imum. The more common freshwater spe- cies were Ceratium hirundinella and Peri- dinium willei. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Summary The tidal fresh region of the James River contained a diverse and abundant phyto- plankton flora. A total of 271 taxa were identified, with the most abundant species chlorophytes (94), diatoms (78), and cyano- bacteria (36). The region was dominated by freshwater algae, but there were also some estuarine diatoms and phytoflagellates re- corded for the site. The recording of estu- arine taxa was probably enhanced by the extensive sampling base, which provided more opportunities for these species to be recorded for this region, e.g., during storm events and periods of low river discharge, etc., over 10.5 years. However, the majority of these species occurred in <2% of the collections. A high degree of representation by estuarine species in this region would not be expected to occur during each monthly collection. There was monthly and annual variability in river discharge at this station. Although the use of mean discharge rates and mean phytoplankton concentrations presented here do not identify close annual relation- ships, a pattern is present. Peak develop- ment of diatoms (the spring pulse) occurred during periods of increased river discharge, while the high populations of chlorophytes, cyanobacteria, autotrophic picoplankton, and euglenophytes were closely associated to periods of reduced flow and more stable water conditions. These findings support re- sults presented by Farrell (1994), Schmidt (1994), and others. Peak periods of phytoplankton abun- dance extended from spring through fall, with different algal categories showing a successional pattern of development. Win- ter was the period of least abundance, bio- mass, and productivity, with low concentra- tions of diatoms, chlorophytes, and cryp- tomonads as the common constituents. The spring bloom of freshwater diatoms, mainly by Skeletonema potamos, was followed by increased concentrations of chlorophytes into early summer. The summer flora con- VOLUME 111, NUMBER 3 tained high concentrations of chlorophytes, cyanobacteria, autotrophic picoplankton, euglenoids, and diatoms. This was the pe- riod for maximum values in phytoplankton and picoplankton abundance, biomass, and productivity. Into fall, the dominance of the summer constituents declined, except for the diatoms, which produced another pulse. This was followed by reduced concentra- tions of total phytoplankton and the auto- trophic picoplankton into winter. Common background taxa to these major components included cryptophytes, euglenophytes, xan- thophytes, and chrysophytes. Acknowledgements This study is a component of the Ches- apeake Bay Phytoplankton Monitoring Pro- gram supported by the Virginia Department of Environmental Quality and the U.S. En- vironmental Protection Agency. The pro- ductivity analysis was conducted by Dr. Kneeland Nesius of Old Dominion Univer- sity. Literature Cited Anderson, G. 1986. Silica, diatoms and a freshwater productivity maximum in Atlantic coastal plain estuaries, Chesapeake Bay.—Estuarine, Coastal, and Shelf Science 22:183-197. Belval, D., J. Campbell, S. Phillips, & C. Bell. 1995. Water-quality characteristics of five tributaries to the Chesapeake Bay at the fall line, Virginia, July 1988 through June 1993. U.S. Geological Survey, Water-Resources Investigation Repport 95-4258, 71 pp. Cloern, J. E. 1987. Turbidity as a control on phyto- plankton biomass and productivity in estuar- ies.—Continental Shelf Research 7:(11/ 12)1367-1381. , A. Alpine, B. Cole, R. Wong, J. Arthur, & M. Ball. 1983. River discharge controls phyto- plankton dynamics in the northern San Francis- co Bay Estuary.—Estuarine, Coastal, and Shelf Science 16:415—429. Farrell, I. 1994. Comparative analysis of the phyto- plankton of fifteen lowland fluvial systems of the River Plate Basin (Argentina).—Hydrobiol- ogia 289:109-117. Filardo, M., & W. Dunstan. 1985. Hydrodynamic con- trol of phytoplankton in low salinity waters of 727 the James River estuary U.S.A.—Estuarine, Coastal, and Shelf Science 21:653—668. Forester, J. W. 1973. The fate of freshwater algae en- tering an estuary. Pp. 387—419 in L. Stevensen and R. Colwell, eds., Estuarine Microbial Ecol- ogy, University of South Carolina Press, Co- lumbia. Haas, L., S. Hastings, & K. Webb. 1981. Phytoplank- ton response to a Stratification-mixing cycle in the York River estuary during late summer. Pp. 619-635 in B. Neilson and L. Cronin, eds., Es- tuaries and Nutrients, Humana Press. Haertel, L., C. Osterberg, H. Curl, & P. Park. 1969. Nutrient and plankton ecology of the Columbia River estuary.—Ecology, 50:962—978. Jackson, R., P. leB. Williams, & I. Joint. 1987. Fresh- water phytoplankton in the low salinity region of the River Tamar Estuary.—Estuarine, Coast- al, and Shelf Science 25:299-311. Jones, R. C. 1991. Spatial and temporal patterns in a cyanobacterial phytoplankton bloom in the tidal freshwater Potomac River, U.S.A.—Verhand- lungen Internationale Vereiningung Limnologie 24:1698—1702. , C. Buchanan, & V. Andrele. 1992. Spatial, seasonal, and interannual patterns in the phy- toplankton communities of a tidal freshwater ecosystem.—Virginia Journal of Science 43:25— 40. Marshall, H. G. 1994. Spatial and temporal diatom re- lationships in the lower James River, Virginia, U.S.A.—Proceedings of the 11th International diatom Symposium, Memoirs of the California Academy of Sciences No. 17:449—457. . 1995. Autotrophic picoplankton distribution and abundance in the Chesapeake Bay, U.S.A.—Marine Nature 4:33—42. , & L. Affronti. 1992. Seasonal phytoplankton development within three rivers in the lower Chesapeake Bay region.—Virginia Journal of Science 43:15—23. , & R. W. Alden. 1990. A comparison of phy- toplankton assemblages and environmental re- lationships in three estuarine rivers of the lower Chesapeake Bay.—Estuaries 13:287—300. , & K. K. Nesius. 1993. Seasonal relationships between phytoplankton composition, abun- dance, and primary productivity in three tidal rivers of the lower Chesapeake Bay.—Journal of the Elisha Mitchell Scientific Society 109: 141-151. Orute, FE I. 1990. Phytoplankton flora in the Warri/ Forcados Estuary of southern Nigeria—Hydro- biologia 208:101—109. Schmidt, A. 1994. Main characteristics for the phyto- plankton of the southern Hungarian section of the River Danube.—Hydrobiologia 289:97— 108. 728 Schuchardt, B., & M. Schirmer. 1991. Phytoplankton maxima in the freshwater reaches of two coastal plain estuaries.—Estuarine, Coastal, and Shelf Science 32:187—206. Smayda, T. 1978. From phytoplankters to biomass. Pp. 273-279 in A. Sournia, ed., Phytoplankton Manual. United Nations Educational, Scientific, and Cultural Organization, Paris. Webb, K. 1988. Comment on “‘Nutrient limitation of phytoplankton growth in brackish coastal ponds” by Caraco, Tamse, Bourtos, and Valiela (1987).’’—Canadian Journal of Fisheries and Aquatic Sciences 45:380—381. Appendix 1 Phytoplankton within the tidal fresh region of the James River. BACILLARIOPHY CEAE Achnanthes sp. Actinocyclus normanii (Gregory) Hustedt Asterionella formosa Hassall Asterionella glacialis Castracane Asterionella gracillima (Hantzsch) Heiberg Aulacoseira distans (Ehrenberg) Simonsen Aulacoseira distans v. humilis (A. Cleve) R. Ross Aulacoseira granulata (Ehrenberg) Simonsen Aulacoseira granulata v. angustissima (O. Miiller) Si- monsen Aulacoseira herzogii (Lemmermann) Simonsen Aulacoseira islandica (O. Miiller) Simonsen Bacillaria paxillifer (Miller) Hendy Chaetoceros sp. Cocconeis sp. Coscinodiscus marginatus Ehrenberg Coscinodiscus sp. Cyclostephanos sp. Cyclostephanos dubius (Fricke in A. Schmidt) E E. Round Cyclotella bodanica Grunow Cyclotella caspia Grunow Cyclotella chaetoceros Lemmermann Cyclotella choctawhatcheeana Prasad Cyclotella comta (Ehrenberg) Kiitzing Cyclotella meneghiniana Kiitzing Cyclotella sp. Cyclotella striata (Kiitzing) Grunow Cylindrotheca closterium (Ehrenberg) Reiman & Lewin Cymbella affinis Kiitzing Cymbella sp. Diatoma sp. Diploneis sp. Ditylum brightwellii (T. West) Grunow Fragilaria capucina Desmaziéres Fragilaria construens (Ehrenberg) Grunow Fragilaria crotonensis Kitton Fragilaria sp. Gyrosigma fasciola (Ehrenberg) Griffith & Henfrey PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gyrosigma littorale (W. Smith) Griffith & Henfrey Gyrosigma sp. Hantzchia sp. Leptocylindrus danicus Cleve Leptocylindrus minimus Gran Licmorphora sp. Melosira moniliformis (O. EF Miller) C. Agardh Melosira varians C. Agardh Meridion circulare (Greville) C. Agardh Navicula radiosa Kiitzing Navicula sp. Nitzschia acicularis (Ktitzing) W. Smith Nitzschia obtusa W. Smith Nitzschia sigma (Kitzing) W. Smith Nitzschia sp. Pinnularia sp. Pleurosigma angulatum (Quekett) W. Smith Pleurosigma elongatum W. Smith Pleurosigma sp. Pseudonitzschia pungens Grunow Rhaphoneis amphiceros (Ehrenberg) Ehrenberg Rhaphoneis surierella (Ehrenberg) Grunow Rhizosolenia delicatula Cleve Skeletonema costatum (Greville) Cleve Skeletonema potamos (Weber) Hasle Stauroneis sp. Staurosirella leptostauron (Ehrenberg) Williams & Round Stephanodiscus hantzschii Grunow Stephanodiscus sp. Surirella elegans Kiitzing Surirella ovata Kiitzing Surirella robusta Ehrenberg Surirella sp. Surirella tenera Gregory Synedra acus Kiitzing Synedra sp. Synedra ulna (Nitzsch) Ehrenberg Tabellaria sp. Thalassionema nitzschioides (Grunow) Grunow Thalassiosira lacustris (Grunow) Hasle & Fryxell Tryblionella punctata W. Smith CHLOROPH YCEAE Acenedesmus acuminatus (Lagerheim) Chodat Actinastrum hantzschii Lagerheim Actinastrum hantzschii v. fluviatile Schréder Ankistrodesmus gracilis (Reinsch) Korschikov Ankistrodesmus falcatus (Corda) Ralfs Ankistrodesmus falcatus v. acicularis (A. Braun) G. S. Smith Ankistrodesmus longissimus (Lemmermann) Wille Carteria fornicata Nygaard Carteria sp. Chlamydomonas sp. Chlamydomonas pertyi Goroshankin Chlorella vulgaris Beyer Closteriopsis acicularis (G. M. Smith) Belcher & Swale VOLUME 111, NUMBER 3 Closteriopsis longissima (Lemmermann) Lemmer- mann Closterium acutum Lyngbye ex Ralfs Closterium sp. Coelastrum microporum Nageli Cosmarium rectangulare Grunow Cosmarium sp. Crucigenia apiculata (Lemmermann) Schmidle Crucigenia crucifera (Wolle) Collins Crucigenia fenestrata (Schmidle) Schmidle Crucigenia tetrapedia (Kirchner) W. ex G.S. West Crucigenia quadrata Morren Crucigenia rectangularis (A. Braun) Gay Crucigenia sp. Dictyosphaerium ehrenbergianum Nageli Dictyosphaerium tetrachotomium Printz Didymocystis planctonica Korschikov Diplocloris hoefleri (Bour) Hindak Endorina cylindrica Korschikov Euastrum gayanum DeToni Franceia elongata Korschikov Hyaloraphidium arcuatum Korschikov Hyaloraphidium contortum Pascher & Korschikov Kirchneriella contorta (Schmidle) Bohlin Kirchneriella lunaris (Kirchner) Moebius Kirchneriella irregularis v. spiralis (Smith) Korschi- kov Kirchneriella obesa (W. West) Schmidle Kirchneriella sp. Koliella longiseta (Vischer) Hindak Micractinium pusillum Fresenius Monoraphidium arcuatum (Korschikov) Hindak Monoraphidium contortum (Thuret) Komarkova-Leg- nerova Monoraphidium grifithii (Berkel) Komarkova-Legne- rova Monoraphidium minutum (Nageli) Komarkova-Leg- nerova Monoraphidium obtusum (Korschikov) Komarkova- Legnerova Monoraphidium pusillum (Printz) Komarkova-Legne- rova Oocystis coronata Lemmermann Oocystis solitaria Wittrock Pediastrum biradiatum Meyen Pediastrum boryanum v. longicornum Reinsch Pediastrum duplex Meyen Pediastrum duplex v. inflata Wolosz Pediastrum simplex (Meyen) Lemmermann Pediastrum tetras (Ehrenberg) Ralfs Phacotus sp. Phacotus lenticularis Ehrenberg Quadrigula lacustris (Chodat) G. M. Smith Quadrigula phitzeri (Schréder) G. M. Smith Scenedesmus acuminatus (Lagerheim) Chodat Scenedesmus armatus Chodat Scenedesmus bicaudatus (Hansgirg) Chodat Scenedesmus bijuga (Turpin) Lagerheim 729 Scenedesmus bijuga vy. alternans (Reinsch) Hansgirg Scenedesmus denticulatus Lagerheim Scenedesmus dimorphus (Turpin) Kiitzing Scenedesmus disciformis (Chodat) Fott & Komarkova Scenedesmus ecornis (Ralfs) Chodat Scenedesmus magnus Meyen Scenedesmus obliquus (Turpin) Kiitzing Scenedesmus opoliensis Richter Scenedesmus quadricauda (Turpin) Brébisson Scenedesmus smithii Lemmermann Scenedesmus sp. Schroederia setigera (Schré6der) Lemmermann Staurastrum americanum (West) G. M. Smith Staurastrum chaetoceras (Schroder) G. S. Smith Staurastrum paradoxum Meyen Staurastrum sp. Staurastrum tetracerum Ralfs Tetraédron arthrodesmiforme (W. West) Woloszynska Tetraédron gracile (Reinsch) Hansgirg Tetraédron minimum (A. Braun) Hansgirg Tetraédron regulare Kitzing Tetraédron sp. Tetraédron triacanthum Korschikov Tetraédron trigonium (Nagel) Hansgirg Tetrastrum elegans Playfair Tetrastrum heteracanthum (Nordstedt) Chodat Tetrastrum staurogeniaeforme (Schroder) Lemmer- mann Treubaria setigera (Archer) D. M. Smith Westella botryoides (W. West) de Wildermann Xanthidium antilopeum Ehrenberg ex Kiitzing CYANOBACTERIA Anabaena affinis Lemmermann Anabaena flos-aquae Brébisson Anabaena reniformis Lemmermann emend. Aptekay Anabaena sp. Anabaena spiroides Klebahn Anabaena solitaria Klebs Aphanizomenon flos-aquae (L.) Ralfs Aphanizomenon issatschenkoi (Ussac.) Proschkina- Lavenko Chroococcus limneticus Lemmermann Chroococcus minor (Kiitzing) Nageli Dactylococcopsis acicularis Lemmermann Dactylococcopsis raphidioides Hansgirg Gomphosphaeria aponina Kitzing Gomphosphaeria fusca Skuja Gomphosphaeria lacustris Chodat Gomphosphaeria naegeliana (Unger) Lemmermann Lyngbya contorta Lemmermann Merismopedia elegans A. Braun Merismopedia glauca (Ehrenberg) Nageli Merismopedia punctata Meyen Merismopedia tenuissima Lemmermann Microcystis aeruginosa Kitzing Microcystis firma (Brébisson & Lemmermann) Schmi- dle Microcystis incerta Lemmermann 730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Microcystis viridis (A. Braun) Lemmermann Microcystis wesenbergii Komarkova Nostoc commune Vaucher Nostoc sp. Oscillatoria agardhii Gomont Oscillatoria limnetica Lemmermann Oscillatoria sp. Phormidium sp. Spirulina major Kiitzing Spirulina sp. Spirulina subsalsa Oersted Synechococcus sp. XANTHOPHYCEAE Botrydiopsis arhiza Borzi Botrydiopsis eriensis Snow Centritractus brunneus Fott Centritractus capilifer Pascher Centritractus globulosus Pascher Characiopsis subulata (A. Braun) Borzi Dichotomococcus curvatus Korschikov Gleobotrys limneticus (G. M. Smith) Pascher Goniochloris pulcherrima Pascher Isthmochloron lobulatum (Nageli.) Skuja Pseudotetraedron neglectum Pascher Tetraedriella spinigera Skuja Tribonema aequale Pascher Tribonema affine G. S. West Tribonema ambiguum Skuja Tribonema monochloron Pascher & Geitler Tribonema pyrenigerum Pascher Tribonema subtilissimum Pascher Tribonema vulgare Pascher EUGLENOPHYCEAE Euglena acus Ehrenberg Euglena ehrenbergii Klebs Euglena gracilis Klebs Euglena oblonga Schmitz Euglena oxyuris Schmarda Euglena tripteris (Dujardin) Klebs Euglena viridis Ehrenberg Phacus lemmermannii (Swirenko) Skvortzow Phacus longicauda (Ehrenberg) Dujardin Phacus suecicus Lemmermann Phacus sp. Strombomonas borysteniensis (Roll) Popova Strombomonas sp. Trachelomonas hispida (Perty) Stein Trachelomonas sp. CRYPTOPHYCEAE Cryptomonas erosa Ehrerberg Cryptomonas curvata Ehrenberg emend. Penard Cryptomonas marssoni Skuja Cryptomonas ovata Ehrenberg Cryptomonas ovata v. curvata (Ehrenberg) Lemmer- mann Rhodomonas minuta Skuja Rhodomonas ovata Ehrenberg CHRYSOPHYCEAE Dictyota fibula Ehrenberg Dinobryon bavaricum Imhof Dinobryon cylindricum Imhof Dinobryon petiolatum Willen Dinobryon sertularia Ehrenberg Dinobryon sociale Ehrenberg Lagynion cystodini Pascher Synura sp. Synura uvella Ehrenberg DINOPHYCEAE Ceratium hirundinella (O. F Miller) Dujardin Ceratium sp. Cochlodinium sp. Gymnodinium sp. Gyrodinium fusiforme Kofoid & Swezy Gyrodinium sp. Heterocapsa triquetra (Ehrenberg) Stein Katodinium rotundatum (Lohmann) Loeblich III Peridinium sp. Peridinium willei Huitfelds-Kaas Prorocentran micans Ehrenberg Prorocentrum minimum (Pavillard) Schiller Protoperidinium sp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):731-734. 1998. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE Applications published in the Bulletin of Zoological Nomenclature The following Applications were published on 31 March 1998 in Vol. 55, Part 1 of the Bulletin of Zoological Nomenclature. Comment or advice on any of these applications is invited for publication in the Bulletin and should be sent to the Executive Secretary (I.C.Z.N.), % The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). Case No. 3011 Strombidium gyrans Stokes, 1887 (currently Strobilidium gyrans) and Stro- bilidium caudatum Kahl, 1932 (Ciliophora, Oligotrichida): proposed conservation of the specific names. Osilinus Philippi, 1847 and Austrocochlea Fischer, 1885 (Mollusca, Gastrop- oda): proposed conservation by the designation of Trochus turbi- natus Born, 1778 as the type species of Osilinus. Androctonus caucasicus Nordmann, 1840 (currently Mesobuthus caucasicus; Arachnida, Scorpiones): proposed conservation of the specific name. Paruroctonus Werner, 1934 (Arachnida, Scorpiones): proposed conservation. Corisa propinqua Fieber, 1860 (currently Glaenocorisa propinqua; Insecta, Heteroptera): proposed conservation of the specific name. Phytobius Dejean, 1835 (Insecta, Coleoptera): proposed conservation. DASYPODIDAE Borner, 1919 (Insecta, Hymenoptera): proposed emenda- tion of spelling to DASYPODAIDAE, so removing the homonymy with DASYPODIDAE Gray, 1821 (Mammalia, Xenarthra). 732 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The following Applications were published on 30 June 1998 in Vol. 55, Part 2 of the Bulletin of Zoological Nomenclature. Comment or advice on any of these ap- plications is invited for publication in the Bulletin and should be sent to the Exec- utive Secretary (I.C.Z.N.), % The Natural History Museum, Cromwell Road, Lon- don SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). Case No. 2956 Campeloma Rafinesque, 1819 (Mollusca, Gastropoda): proposed conserva- tion 3008 Euchilus Sandberger, 1870 and Stalioa Brusina, 1870 (Mollusca, Gastropo- da): proposed designation of Bithinia deschiensiana Deshayes, 1862 and Paludina demarestii Prévost, 1821 as the respective type spe- cies, with the conservation of Bania Brusina, 1896 Holospira Martens, 1860 (Mollusca, Gastropoda): proposed designation of Cylindrella goldfussi Menke, 1847 as the type species Thamnotettix nigropictus Stal, 1870 (currently Nephotettix nigropictus; In- secta, Homoptera): proposed conservation of the specific name Cicada clavicornis Fabricius, 1794 (currently Asiraca clavicornis; Insecta, Homoptera): proposed conservation of the specific name Musca rosae Fabricius, 1794 (currently Psila or Chamaepsila rosae; Insecta, Diptera): proposed conservation of the specific name Iguanodon Mantell, 1825 (Reptilia, Ornithischia): proposed designation of Iguanodon bernissartensis Boulenger in Beneden, 1881 as the type species, and proposed designation of a lectotype VOLUME 111, NUMBER 3 733 Opinions published in the Bulletin of Zoological Nomenclature The following Opinions were published on 31 March 1998 in Vol. 55, Part 1 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History Mu- seum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). Opinion No. 1886. Plumularia Lamarck, 1816 (Cnidaria, Hydrozoa): conserved by the desig- nation of Sertularia setacea Linnaeus, 1758 as the type species. 1887. Arca pectunculoides Scacchi, 1834 and A. philippiana Nyst, 1848 (currently Bathyarca pectunculoides and B. philippiana, Mollusca, Bivalvia): specific names conserved. Lirobarleeia Ponder, 1983 (Mollusca, Gastropoda): Alvania nigrescens Bartsch & Rehder, 1939 designated as the type species. Parapronoe crustulum Claus, 1879 (Crustacea, Amphipoda): specific name conserved. Scarabaeus rufus Moll, 1782 (currently Aphodius rufus), Scarabaeus rufus Fabricius, 1792 (currently Aegialia rufa) and Scarabaeus foetidus Herbst, 1783 (currently Aphodius foetidus) (Insecta, Coleoptera): specific names conserved. Crenitis Bedel, 1881, Georissus Latreille, 1809 and Oosternum Sharp, 1882 (Insecta, Coleoptera): conserved. Alcyonidium mytili Dalyell, 1848 (Bryozoa): neotype replaced. Bombycilla cedrorum Vieillot, [1808] and Troglodytes aedon Vieillot, [1809] (Aves, Passeriformes): specific names conserved. Regnum Animale ..., Ed. 2 (M.J. Brisson 1762): rejected for nomenclatural purposes, with the conservation of the mammalian generic names Philander (Marsupialia), Pteropus (Chiroptera), Glis, Cuniculus and Hydrochoerus (Rodentia), Meles, Lutra and Hyaena (Carnivora), Tapirus (Perissodactyla), Tragulus and Giraffa (Artiodactyla). 734 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The following Opinions were published on 30 June 1998 in Vol. 55, Part 2 of the Bulletin of Zoological Nomenclature. Copies of these Opinions can be obtained free of charge from the Executive Secretary, I.C.Z.N., % The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk). Opinion No. 1895. Riisea and riisei Duchassaing & Michelotti, 1860 (Cnidaria, Anthozoa): con- served as the correct original spellings of generic and specific names based on the surname Riise 1896. Galba Schrank, 1803 (Mollusca, Gastropoda): Buccinum truncatulum Mill- er, 1774 designated as the type species 1897. Glomeris Latreille, 1802 (Diplopoda), Armadillo Latreille, 1802, Armadil- lidium Brandt in Brandt & Ratzeburg, [1831] and Armadillo vulgaris Latreille, 1804 (currently Armadillidium vulgare)(Crustacea, Isopo- da): generic and specific names conserved Metaphycus Mercet, 1917 (Insecta, Hymenoptera): given precedence over Aenasiodea Girault, 1911 Meristella Hall, 1859 (Brachiopoda): Atrypa laevis Vanuxem, 1842 desig- nated as the type species Trematospira Hall, 1859 (Brachiopoda): Spirifer multistriatus Hall, 1857 designated as the type species Gladiolites geinitzianus Barrande, 1850 (currently Retiolites geinitzianus; Graptolithina): lectotype replaced by a neotype PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(3):735-736. 1998. BIOLOGICAL SOCIETY OF WASHINGTON 125th Annual Meeting, 7 May 1998 President Stephen D. Cairns called the meeting to order at 11:00 a.m. in the Waldo Schmitt Room, National Museum of Natu- ral History. Other council members present: Richard P. Vari (President-elect), Carole C. Baldwin (Secretary), T. Chad Walter (Trea- surer), Alfred L. Gardner, Susan L. Jewett, and Rafael Lemaitre (Elected Council), C. Brian Robbins (Editor), Storrs L. Olson (Custodian of Publications), Richard C. Banks and Austin B. Williams (Finance Committee), and Bruce B. Collette and Charles Handley (past BSW presidents). Elected Council not in attendance: John A. Fornshell, Diana Lipscomb, and James N. Norris. Following approval of the minutes of the 124th Annual Meeting of the Society, Pres- ident Cairns announced that Jon L. Noren- burg resigned as Associate Editor for In- vertebrates and was replaced by Stephen L. Gardiner from Bryn Mawr College. On be- half of the Society, President cairns thanked Norenburg for his 4% years of service. He further noted that the appointment of Gar- diner represents the first time a scientist from outside the Smithsonian community has served as an associate editor for the Proceedings; the new arrangement appears to be successful and may set a new prece- dent for future editorial appointments. Treasurer Chad Walter summarized So- ciety finances for the period between 1 Jan- uary and 31 December 1997 (Table 1). To- tal income was $75,333.27 ($40,248.25 from publication charges, $22,041.00 from dues and subscriptions, and $13,044.02 from sales of back issues and interest on Society accounts). Expenditures totaled $75,887.08 ($66,771.14 for publication costs, $8732.43 for management costs, and $383.51 for bank charges). Net change for the period was —$553.81; this is not a real loss but a relative value based on the spe- cific range of time considered. The treasurer noted that interest income increased signif- icantly in 1997 because of new investment strategies planned and implemented by the Finance Committee and a favorable stock market. A discussion about the purpose and ideal maximum value of the Society’s En- dowment Fund was initiated by NMNH Cu- rator Brian Kensley, and President Cairns asked the Finance Committee to establish an approximate monetary figure that could serve as the Society’s goal for the Fund. Richard Banks then gave the Finance Committee Report. He noted that the Fi- nance Committee will begin meeting quar- terly rather than annually with the Treasurer in efforts to refine procedures related to es- tablishing a meaningful annual budget for the Society. As an example of budgetary problems, Banks pointed out that the soci- ety paid ca. $10,000 in unbudgeted publi- cation costs for Bulletin 9. Although those costs were recovered in 1998, actual ex- penses in 1997 greatly exceeded the amount approved in the proposed 1997 budget. President Cairns thanked Banks, Austin Williams and Chad Walter for their help and hard work in maintaining Society fi- nances. He then called upon Brian Robbins — for the Editor’s Report. Four issues of Vol- ume 110 of the Proceedings were published comprising 70 papers and 692 pages. Ad- ditionally, a 149-page Bulletin (No. 9) was published in August. Submissions to the Proceedings increased significantly in 1997 (100 vs. 80 in 1996, 87 in 1995), but as of 1 May 1998 there were 30 submissions, down from 41 in 1997. Robbins noted that there is no backlog for accepted papers. The President then asked Richard Banks 736 Table 1.—Summary financial statement for 1997. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Total assets Endowment fund General fund ASSETS: JANUARY 1, 1997 TOTAL RECEIPTS FOR 1997 TOTAL DISBURSEMENTS FOR 1997 ASSETS: DECEMBER 31, 1997 NET CHANGES IN FUNDS 27,484.69 79,349.34 106,834.03 63,125.28 12,207.99 75,333.27 75,887.08 —.— 75,887.08 14,722.89 ONS) 7/-333)"* 106,280.22 (— 12,761.80) 12,207.99 (—553.81) * The income from publication inventory ($4244) and remainder after opening Merrill Lynch Account ($2330.56) are part of the Endowment Fund, but were left in the General Fund (Riggs account) for cash flow maintenance. to read the proposed changes to Article 8 of the Bylaws, which the Council had agreed in 1997 to amend. By unanimous vote, Article 8 of the Bylaws will read as follows: A Endowment Fund. There shall be an Endowment Fund which shali consist of contributions from members, miscella- neous gifts, and surplus funds from opera- tions. At the discretion of the Council, the principal of this fund may be used (invaded for use) in publishing the Society’s journal or for the general operations of the Society. At the discretion of the Council, the prin- cipal of this Fund may also be used in the publication of symposia, monographic stud- ies, or other special publications; however, such a decision must be reached only dur- ing a regularly scheduled meeting of the Council. Storrs Olson, Custodian of Publications, then announced that the Society had sold two complete sets of back issues of the Pro- ceedings and that profits from those sales exceeded labor, shipping, and advertising costs. Olson noted that four complete sets of back issues remain but that no additional sets will be compiled because reserves of many issues are depleted. Instead, Olson proposed that the remaining back issues that were published as separates (rather than as quarterly issues) be compiled into sub- ject sets, e.g., on fishes, birds, crustaceans, etc., and be advertised and sold this way. There was general agreement that subject sets would be marketable, and Olson agreed to initiate and oversee the sorting. The President thanked Janet W. Reid and Michael D. Carleton for serving on the Nominating Committee for the 1998 elec- tion of officers. Election results, announced by the Secretary, are as follows: President- elect—Brian E Kensley; Secretary—Carole C. Baldwin; Treasurer—T. Chad Walter; Elected Council—Michael D. Carleton, W. Duane Hope, Susan L. Jewett, Rafael Le- maitre, Roy W. McDiarmid, James N. Nor- ris. President Cairns then turned the meet- ing over to incoming President Richard P. Vari who thanked Cairns for his service. Vari adjourned the meeting at 11:34 a.m. Respectfully submitted, Carole C. Baldwin Secretary INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with an Abstract in an alternate language when appropriate. Submission of manuscripts.—Submit three copies of each manuscript in the style of the Proceedings to the Editor, complete with tables, figure captions, and figures (plus originals of the illustrations). Mail directly to: Editor, Proceedings of the Biological Society of Wash- ington, National Museum of Natural History NHB-108, Smithsonian Institution, Washington, D.C. 20560. (Do not submit manuscripts to an associate editor). Presentation.—Clarity of presentation, and requirements of taxonomic and nomenclatural procedures necessitate reasonable consistency in the organization of papers. Telegraphic style is recommended for descriptions and diagnoses. The style for the Proceedings is described in “GUIDELINES FOR MANUSCRIPTS for Publications of the BIOLOGICAL SOCIETY OF WASHINGTON,’ a supplement to Volume 103, number 1, March 1990. Authors are encour- aged to consult this article before manuscript preparation. Copies of the article are available from the editor or any associate editor. The establishment of new taxa must conform with the requirements of appropriate interna- tional codes of nomenclature. Decisions of the editor about style also are guided by the General Recommendations (Appendix E) of the International Code of Zoological Nomenclature. When appropriate, accounts of new taxa must cite a type specimen deposited in an institutional col- lection. Review.—One of the Society’s aims is to give its members an opportunity for prompt pub- lication of their shorter contributions. Manuscripts are reviewed by a board of Associate Editors and appropriate referees. Proofs.—Authors will receive first proofs and original manuscript for correction and ap- proval. Both must be returned within 48 hours to the Editor. Reprint orders are taken with returned proofs. Publication charges.—Authors are required to pay full costs of figures, tables, changes in proofs ($3.00 per change or revision), and reprints. Authors are also asked to assume costs of page-charges. The Society, on request, will subsidize a limited number of contributions per volume. Payment of full costs will facilitate speedy publication. Costs.—Printed pages @ $65.00, figures @ $10.00, tabular material @ $3.00 per printed inch per column. One ms. page = approximately 0.4 printed page. CONTENTS Horton H. Hobbs, Jr. (29 March 1914—22 March 1994). Biographical notes Karen Reed and Raymond B. Manning Franconictis (Mammalia: Carnivora) from the Late Oligocene of eastern Kazakstan Spencer G. Lucas, Robert J. Emry, and Pyruza A. Tleuberdina Diagnoses of hybrid hummingbirds (Aves: Trochilidae). 6. An intergeneric hybrid, Aglaiocer- cus kingi X Metallura tyrianthina, from Venezuela Gary R. Graves A new species of Alsodes (Amphibia: Anura: Leptodactylidae) from southern Chile J. Ramon Formas, César Cuevas, and José Nunez Helicoprion nevadensis (Wheeler, 1939) from the Pennsylvanian—Permian Antler Peak Lime- stone, Lander County, Nevada (Pisces: Selachii: Helicoprionidae) Rex Alan Hanger and Ellen E. Strong Pentamera rigida and P. pediparva, two new species of sea cucumber from the west coast of North America (Echinodermata: Holothuroidea) Philip Lambert The Anacroneuria of Costa Rica and Panama (Insecta: Plecoptera: Perlidae) Bill P. Stark Studies in aquatic insects XIV: Description of eight new species of Ochrotrichia Mosley (Tri- choptera: Hydroptilidae), from Costa Rica Joaquin Bueno-Soria and Ralph Holzenthal A new species of the genus Gastrosaccus (Crustacea: Mysidacea: Mysidae) from Oman Masaaki Murano and Anton McLachlan A new species of amphipod (Crustacea: Amphipoda: Lysianassoidea) from the Pacific Coast of North America Ann Dalkey Metatiron bonaerensis, a new species (Crustacea: Amphipoda: Synopiidae) from the southwest Atlantic Gloria M. Alonso de Pina A new genus and species of “‘goneplacid-like’’ brachyuran crab (Crustacea: Decapoda) from the Gulf of California, Mexico,and a proposal for the use of the family Pseudorhombildae Alcock, 1900 Michel E. Hendrickx A new crayfish of the genus Orconectes from the Blood River drainage of western Kentucky and Tennessee (Decapoda: Cambaridae) Christopher A. Taylor and Mark H. Sabaj Two new species of Erugosguilla from the Indo-West Pacific (Crustacea: Stomatopoda: Squil- lidae) Shane T. Ahyong and Raymond B. Manning Two new species of Macrobiotidae (Tardigrada: Eutardigrada) from the United States of Amer- ica, and some taxonomic considerations of the genus Murrayon Roberto Guidetti The genus Perinereis (Polychaeta: Nereididae) from Mexican littoral waters, including the description of three new species and the redescriptions of P. anderssoni and P. elenacasoae Jestis Angel de Leén Gonzalez and Vivianne Solis-Weiss Streblospio gynobranchiata, a new spionid polychaete species (Annelida: Polychaeta) from Florida and the Gulf of Mexico with an analysis of phylogenetic relationships within the genus Streblospio Stanley A. Rice and Lisa A. Levin Capitellids (Polychaeta: Capitellidae) from the continental shelf of the Gulf of California, México, with the description of a new species, Notomastus angelicae Pablo Hernandez-Alcantara and Vivianne Solis-Weiss Phytoplankton composition within the tidal freshwater region of the James River, Virginia Harold G. Marshall and Lubomira Burchardt International Commission on Zoological Nomenclature Biological Society of Washington: 125th Annual Meeting 473 674 694 708 720 731 135 : PROCEEDINGS BOF THE ~ BIOLOGICAL SOCIETY OF _ WASHINGTON VOLUME 111 NUMBER 4 23 DECEMBER 1998 ISSN 0006-324X THE BIOLOGICAL SOCIETY OF WASHINGTON 1998-1999 Officers President: Richard P. Vari Secretary: Carole C. Baldwin President-elect: Brian F. Kensley ; Treasurer: TY. Chad Walter Elected Council Michael D. Carleton Rafael Lemaitre W. Duane Hope Roy W. McDiarmid Susan L. Jewett James N. Norris Custodian of Publications: Storrs L. Olson PROCEEDINGS Editor: C. Brian Robbins Associate Editors Classical Languages: Frederick M. Bayer Invertebrates: Stephen L. Gardiner Plants: David B. Lellinger Frank D. Ferrari Insects: Wayne N. Mathis Rafael Lemaitre Vertebrates: Gary R. Graves Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $25.00 (for USA and non-USA addresses) include subscription to the Pro- ceedings of the Biological Society of Washington. Annual dues are payable on or before January 1 of each year. Renewals received after January 1 must include a penalty charge of $3.00 for reinstatement. Library subscriptions to the Proceedings are: $40.00 for USA and non-USA addresses. Non-USA members or subscribers may pay an additional $25.00 to receive the Proceedings by Air Mail. The Proceedings of the Biological Society of Washington (USPS 404-750) is issued quarterly. Back issues of the Proceedings and the Bulletin of the Biological Society of Washington (issued sporadically) are available. Correspondence dealing with membership and subscriptions should be sent to: BIOLOGICAL SOCIETY OF WASHINGTON P.O. BOX 1897 LAWRENCE, KANSAS 66044, U.S.A. Payment for membership is accepted in US dollars (cash or postal money order), checks on US banks, or MASTERCARD or VISA credit cards. Manuscripts, corrected proofs, and editorial questions should be sent to: EDITOR BIOLOGICAL SOCIETY OF WASHINGTON NATIONAL MUSEUM OF NATURAL HISTORY WASHINGTON, D.C. 20560, U.S.A. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Periodicals postage paid at Washington, D.C., and additional mailing office. POSTMASTER: Send address changes to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON, P.O. Box 1897, Lawrence, Kansas 66044. This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):737-773. 1998. Sponges, genus Mycale (Poecilosclerida: Demospongiae: Porifera), from a Caribbean mangrove and comments on subgeneric classification Eduardo Hajdu and Klaus Riitzler (EH) Institute of Systematics and Population Biology, University of Amsterdam, P.O. Box 94766 1090-GT, Amsterdam, The Netherlands; Present address: Departamento de Invertebrados, Museu Nacional, Universidade Federal do Rio de Janeiro, Quinta da Boa Vista, s/n 20940-040, Rio de Janeiro, RJ; and Centro de Biologia Marinha, Universidade de Sao Paulo, Sao Sebastiao, SP, Brazil; (KR) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract.—Eight species of Mycale Gray (Mycalidae, Poecilosclerida, De- mospongiae) are described from marine mangroves on the barrier reef of Be- lize, Central American Caribbean. Two are new: Mycale (Aegogropila?) car- migropila and M. (Ae.) citrina. Other species found are M. (Ae.) arndti, M. (Arenochalina) laxissima, M. (Carmia) magnirhaphidifera, M. (C.) microsig- matosa, M. (Mycale) laevis and M. (Paresperella) sp. A key to the 17 recog- nized Caribbean species of Mycale is provided. Ectosomal skeletal patterns currently used as diagnostic characters for subgenera of Mycale may be inad- equate for phylogenetic analysis, but reliable alternative congruent traits have not yet been identified to replace these. More than 150 species of Mycale Gray have been described worldwide (Doumenc & Lévi 1987), with representatives in most marine habitats. They are common in. both polar and tropical seas, and have been re- ported from intertidal pools abyssal depths (Hartman 1982). Contemporary mono- graphs have added great numbers of new species (Lévi 1963, van Soest 1984, Berg- quist & Fromont 1988, Hajdu & Desquey- roux-Fatindez 1994), indicating that many more undescribed taxa are yet to be found. Our study of the rich marine mangrove eco- system of Belize (Riitzler & Feller 1988, 1996; de Weerdt et al. 1991) is no exception and has revealed two new species of Myc- ale. Recent findings of metabolites with pharmacological potential from species of Mycale (e.g., Capon & Macleod 1987; Per- ry et al. 1988, 1990; Fusetani et al. 1989; Corriero et al. 1989; Butler et al. 1991; Northcote et al. 1991; Hori et al. 1993) have strengthened the need for a better as- sessment of the genus’ biodiversity, for a stable system of classification, and for bet- ter descriptions to differentiate between al- lopatric sibling species. It has been convincingly argued that de- scriptions of Mycale are often unreliable in respect to noting size categories of micro- scleres (Hentschel 1913, Doumenc & Lévi 1987). Accordingly, special attention was here paid to this important characteristic. Details of microsclere shape too were long considered to be useful characters because of their low adaptive value (Ridley & Den- dy 1887, Dendy 1921, Hajdu et al. 1994a, Hajdu & Desqueyroux-Fatindez 1994). This view is adopted here and supported by our scanning electron micrography. Materials and Methods Sponges were collected by the authors and Kathleen P. Smith during several sur- Mexico Caribbean Sea Twin Cays sy ens 3: SENG, oe Blue Ground pe gamer ig ig ig “8° Carrie Bow Cay ‘Curlew Bank Fig. 1. 88°08.89'W, Blueground Range. veys of mangroves in the vicinity of Carrie Bow Cay, Belize (16°48’N, 88°05'’W) (Fig. 1). Specimens are deposited in the sponge collection of the National Museum of Nat- ural History, Washington, subsamples were donated to the Museu Nacional, Universi- dade Federal do Rio de Janeiro, Brazil. Schizotypes of the two new species were also deposited in The Natural History Mu- seum, London, Queensland Museum, Bris- bane, and Zodlogisch Museum Amsterdam. Most anatomical preparations were made according to Riitzler (1978) and Hajdu (1994). Additionally, skeletons for study by :Q south Water Cay PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lair ! “Grouper... * Gardens : Map of collecting area on the barrier-reef platform of Belize. Geographical coordinates for principal localities are 16°48.18’N, 88°04.93’W, Carrie Bow Cay; 16°49.95'N, 88°06.34'W, Twin Cays; and 16°48.55’N, scanning electron microscopy (SEM) were prepared in one of the two following ways: Samples in ethanol were washed of debris with several jets of ethanol, air dried in an oven, and mounted on SEM stubs by ap- plying a thin layer of Entellan (Merck); samples from ethanol were washed in warm water (ca. | min), immersed in a saturated solution of soda, and heated to 50—70°C (30 min). Then they were again washed in warm water (1 min) and etched in 20% HNO, at room temperature. After a few minutes under observation, samples were rinsed with warm-water jets (1 min) and a VOLUME 111, NUMBER 4 few drops of hydrogen peroxide (H,O,) so- lution added to the last change of water. Af- ter 30 min at 5O—70°C the samples were rinsed in warm water (1 min), transferred to ethanol (96%, 30 min), air-dried in an oven, and mounted on SEM stubs. Abbreviations used are: BMNH (The Natural History Museum, London), INV- POR (Invemar-Porifera Collection, Univer- sidad Nacional de Colombia, Santa Marta), MCZ (Museum of Comparative Zoology, Harvard University, Cambridge), MNHN (Muséum National d’Histoire Naturelle, Paris), MNRJ (Museu Nacional, Universi- dade Federal do Rio de Janeiro), MSNG (Museo Civico di Storia Naturale di Gen- ova), MUT (Museo della Universita di To- rino), NNM (National Natuurhistorisch Mu- seum, Leiden), QM (Queensland Museum, Brisbane), UFRJPOR (Universidade Fed- eral do Rio de Janeiro, Porifera collection), USNM (National Museum of Natural His- tory, Washington), and ZMA POR (Zod6lo- gisch Museum Amsterdam, Porifera collec- tion). Systematic Descriptions Order Poecilosclerida Topsent, 1928 Suborder Mycalina Hajdu et al., 1994 Family Mycalidae Lundbeck, 1905 Genus Mycale Gray, 1867; sensu Hajdu & Desqueyroux-Fatindez 1994 Diagnosis.—Mycalidae with skeleton of subtylostyles and palmate anisochelas; ad- ditional microscleres may include sigmas, toxas, micracanthoxeas, raphides, unguifer- ate anisochelas, and palmate isochelas. Subgenus Aegogropila Gray, 1867 Diagnosis.—Mycale with tangential ec- tosomal skeleton of reticulate spicule tracts (often easily peeled off), without serrated sigmas. Type species: Aegogropila varians Gray, 1867 (=M. [Ae.] contarenii [von Martens, 1824, as Spongia contarenii.]). Remarks.—Bergquist & Fromont (1988) in quoting Halichondria aegagropila John- 739 ston, 1842 (misspelled as H. aegogropila) as a type-species of Aegogropila (Thiele 1903) overlooked Gray’s (1867) name, Ae- gogropila varians. Because Ae. varians was probably a replacement name in order to avoid tautology, although not clearly stated so by Gray (1867), we choose the figured specimen of H. aegagropila in Johnston (1842, BMNH 1847.9.7.39) as lectotype of Ae. varians. In this way, both species be- come objective synonyms. This point was made clear by L. B. Holthuis (in lit.). The plasticity reported here for the pres- ence of ectosomal reticulation in Mycale (Ae. ?) carmigropila new species is taken as indication that some representatives of Car- mia (see below) are more closely related to Aegogropila than to other species of Car- mia, suggesting paraphyly of the former subgenus and polyphyly of the latter. Ad- ditionally, some M. (Mycale) species may have their confused tangential ectosomal skeleton developed so thinly as to make spicules strewn at random appear reticulat- ed to the casual observer (cf. M. [M.] thielei Hajdu & Desqueyroux-Fatindez 1994). Species like this have been assigned to Ae- gogropila in the past, for instance M. (M.) flagelliformis (Bergquist & Fromont, 1988) which has pore-grooves and a confused ec- tosomal skeleton. It becomes apparent that the monophyletic status of Aegogropila may not hold up in a thorough revision of all species. Such a revision is not yet pos- sible, given that collections are dispersed, many new taxa are still being discovered, and, more importantly, no other congruent characters are yet apparent to replace the existing system. Mycale (Aegogropila) arndti van Soest, 1984 Figs. 2, 3, 17a; TVable 1 Esperia macilenta.—Carter 1871:276, pl. 17, fig. 8; not Hymeniacidon macilenta Bowerbank, 1866 (=Mycale (Carmia) macilenta). Mycale macilenta.—Arndt 1927:143, in PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. b, Detail of surface reticulation showing sigmas-I disposed around the bundles of megascleres, and disposition of rosettes (R) in the corners of the meshes; scale equals 100 wm; c, d, Fusiform subtylostyles and all four categories of anisochelas (I-IV); scale equals 100 wm. (a—c, USNM 39329; d, holotype ZMA POR 3675.) part (not the figured specimen); not AHy- meniacidon macilenta Bowerbank, 1866 (=Mycale (Carmia) macilenta). M. (Aegogropila) arndti van Soest, 1984: 19, pl. Il, fig. 1. Zea 1987:140. Material studied.—ZMA POR 3675, ho- lotype, Curacao; ZMA POR 3842, paratype, Florida. USNM 43032, Belize, Twin Cays, Batfish Point, <1 m, coll. K. Riitzler, 27 Apr 1989; USNM 39329 (MNRJ 630), Be- lize, Blue Ground Range, | m, coll. E. Haj- du & K. Smith, 12 Aug 1993; USNM 47871, Blueground Range, on red-man- grove stilt root, <1 m, coll. K. Smith, Nov 1996. Diagnosis.—Bluish, crustose Aegogro- pila with sigmas (three categories) and tox- as; four categories of anisochelas, with cat- egories II and IV having exceptionally long frontal alae (alae of head and foot nearly fused), covering the entire shaft in frontal view. Mycale arndti, skeleton: a, Peeled-off surface showing ectosomal reticulation; scale equals 500 jm; Description.—Macroscopic appearance: Grayish blue to gray crust or cushion with oscules often raised like small chimmneys; non-slimy when rubbed. Preserved material is composed of seven main fragments, the largest is 55 mm long, 30 mm wide, and up to 4 mm thick. The color in alcohol is drab or pink, varying from light pink to almost violet. The fragments have firm consistency and microhispid surface texture. The ecto- some has sand grains embedded in the sur- face and is easily peeled off. No oscules are apparent. Skeleton: The ectosomal skeleton is a tangential reticulation of 120 wm thick sub- tylostyle bundles (Fig. 2a), with no appar- ent cementing spongin. Meshes are mostly triangular and up to 400 X 200 pm in di- ameter. Microscleres occur in abundance and may be playing the structural role of holding the reticulation intact. Sigmas I (as many as 10 per linear mm) and rosettes VOLUME 111, NUMBER 4 (135 pm, 6 per mm) of anisochelas I can be seen surrounding the megasclere bundles (Fig. 2b). These and other microscleres, oc- cur in great numbers inside the meshes. The choanosome area appears light brown in transmitted light, with dark-brown blotches of fibrous spongin. The spiculation here is not very dense and no clear pattern is apparent which may be due to the slightly macerated condition of the specimen. Mi- croscleres are scattered throughout this area. Spicules (Figs. 2c, d, 3; Table 1): Sub- tylostyles fusiform, gradually tapering to a point, generally slightly bent in upper third and thickest at mid length, with long neck and eliptical head. Anisochelas I stout, with head length 50% of total length, foot in side view at about 110° angle to shaft; frontal alae of foot may bear denticulation on top (Fig. 3e). Anisochelas II similar to aniso- chelas I but even stouter, with head length 70% of total length, shaft not visible in frontal view, lateral alae of head arcuate. Anisochelas III slender, with head length 70% of total length, shaft slightly bent at the end of the lateral alae of the head. An- isochelas IV very slender, with both frontal alae prolonged into thin, digitiform pro- cesses that sometimes cross each other. Sig- mas I stout, abruptly bent into very sharp hooks, with almost straight inner faces. Sig- mas II slender, with gradual curve and sharp hooks. Sigmas III similar to sigmas II but smaller. Toxas very gently bent (more pronounced in short forms). Ecology.—Associated with mangrove and shallow seagrass bottoms in bays and lagoons. Distribution.—Florida, St. Vincent (Vir- gin Islands), Curagao, Colombia, Belize. Remarks.—Only one of this species specimen was found, suggesting that it may be uncommon in the survey area. Details of spicule morphology and presence of four types of anisochelas agree well with the ho- lotype (Figs. 2d, 3b—d). It should be emphasized that proper as- sessment of microsclere categories is essen- 741 tial if a morphological-phylogenetic classi- fication of Mycale is to be achieved. In many taxonomic descriptions, one or the other microsclere category is often over- looked or lost to lumping decisions, as in the present species where anisochelas-cat- egory III was not recognized by previous authors (van Soest 1984, Zea 1987). Con- sequently, anisochelas II of M. arndti should be described as arcuate because the lateral alae of the head are partly separated from the shaft (Hajdu et al. 1994). The stout sigmas I are comparable to those of Esperella simonis as figured by Ridley & Dendy (1887) and of M. quadri- partita Boury-Esnault (Hajdu & Desquey- roux-Fatndez 1994), and may be homolo- gous to the diancistras of Hamacantha Gray (Hajdu 1994). This similarity is based on the straight inner face and the abrupt taper- ing of the hooks in this type of sigma and is enhanced by a faintly marked notch at mid-length of some of these spicules (Fig. 3h). Toxas show a remarkable range of size but size-frequency analysis of 100 toxas in the Belizean specimen failed to reveal dis- tinct categories. Mycale (Aegogropila?) carmigropila, new species Figs. 4, 17b; Table 2 Type material.—Holotype, USNM 34560, Twin Cays, Sponge Haven, on roots of Rhizophora, 1 m, coll. E. Hajdu & K. Smith, 17 Aug 1993. Paratypes: USNM 42997, Belize, Twin Cays, Batfish Point, on red-mangrove stilt root and peat bank, <1 m, coll. K. Riitzler and K. Smith, 5 May 1986; USNM 43048, Belize, Twin Cays, Hidden Creek, on red-mangrove stilt root, <1 m, coll. K. Riitzler, 8 May 1987; USNM 43047, Belize, Twin Cays, Hidden Creek, encrusting a cluster of algae, genus Hali- meda, <1 m, coll. K. Smith, 8 May 1987; USNM 47870, Belize, Twin Cays, Sponge Haven, peat bank with algae, genus Hali- meda, 1 m, coll. K. Smith, 5 Jul 1990; USNM 34561, Twin Cays, Sponge Haven, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Mycale arndti, spicules: a, Face view of anisochela-I; scale equals 10 1m; b, Face view of anisochela- II showing robustness and extremely long frontal ala of head, almost fusing with foot; note deeply concave top portion of frontal ala of the foot (arrow); scale equals 10 wm; c, Oblique frontal view of anisochela-III; scale equals 5 m; d, Profile view of anisochela-[V showing thin digitiform prolongations from both frontal alae (arrows) almost fusing; scale equals 5 tm. e, Detail of foot of anisochela-I in Fig. 3a showing tridentate (t) top VOLUME 111, NUMBER 4 on roots of Rhizophora, 1 m, coll. E. Hajdu & K. Smith, 17 Aug 1993; USNM 34587, Twin Cays, Sponge Haven, on roots of Rhi- zophora, 1 m, coll. E. Hajdu & K. Smith, 17 Aug 1993; USNM 38768, Twin Cays, Sponge Haven, on Halimeda, 1 m, coll. E. Hajdu & K. Smith, 17 Aug 1993. Schizo- types from holotype, BMNH 1994.3.1.1, QM G313152, MNRJ 631, ZMA POR 10708; fragments of paratypes, MNRJ 632, 633, 634. Diagnosis.—Intensely blue to greenish, encrusting Mycale (Aegogropila?) with three categories of anisochelas and one type of small (<70 pm) sigmas. Description.—Macroscopic appearance: Specimens were encrusting (=20 cm’) on roots of Rhizophora, or occurred intermin- gled with Halimeda. In the first case they were cobalt-blue, while in the second they appeared very light green. Other specimens observed alive were blue to violet, turning tan when dying. Subectosomal canals. con- verging on oscules in a star-like pattern are visible in live specimens. The consistency is very soft, slimy, and fragile, the texture rather smooth. Skeleton: The ectosomal skeleton is a polymorphic feature in this species. Two specimens (USNM 34560, 34587) have a tangential reticulation of subtylostyles, sin- gle or in 50 pm thick bundles, with large amounts of cementing spongin. Meshes are mostly triangular (250 < 150 pm). Up to 12 pores (40 X 20 to 80 X 50 pm in di- ameter) of the aquiferous system can be seen within a single mesh. A few rosettes of anisochelas I (<=110 pm in diameter), and sigmas I occur scattered among the meshes, frequently around the bundles. The 60 wm), and copious mucus production when han- dled. Description.—Macroscopic appearance: Thick incrustations (5 mm; 30 cm?) with protruding oscules encircled by a thin mem- brane. Extremely soft and fragile, and re- leasing copious amounts of mucus upon handling. Live-color is lemon yellow to light orange, turning pale yellow in alcohol. Numerous, pronounced subectosomal chan- nels may cover most of the sponge surface and converge toward the oscules. Most specimens contained embryos during Au- gust. Skeleton (Fig. 5): The ectosomal skele- ton peels off easily and is made of a re- markably pure (no scattered megascleres) tangential reticulation of bundled (=85 wm thick) subtylostyles (Fig. 5a), mostly form- ing triangular meshes (660 X 400 pm), with no apparent cementing spongin. Ro- settes of anisochelas-I (130 wm in diameter) occur in places (Fig. 5b, c). Sigmas are very abundant, scattered among the meshes (up to 35 per mesh), and along the strands (30 per mm; Fig. 5d); they seem to play an im- portant structural role in the skeleton. The choanosome is crowded with dark- brown granules (about 40 wm in diameter), presumably of mangrove peat, which ham- per observation of the skeletal architecture. The underlying pattern seems to be densely spicular, with crisscrossing megasclere bun- dles (150 wm thick) and abundant micro- scleres. Spicules (Fig. 6, Table 3): Subtylostyles straight, slender and slightly fusiform, with poorly marked eliptic head. Anisochelas I with straight shaft, head 50% of total length. Anisochelas-II with shaft bent at head, head 50—60% of total length; frontal alae of foot with short, digitiform processes on top (Fig. 6c). Anisochelas-III very slen- der, with shaft gradually curved (profile VOLUME 111, NUMBER 4 Fig. 5. 747 Sel me ’ ee =e a | ie Mycale citrina, skeleton: a, Peeled-off surface showing ectosomal reticulation; scale equals 500 ym; b, Detail of surface reticulation showing sigmas disposed abundantly around the bundles of megascleres, and rosette of anisochelas-I; scale equals 200 wm; c, Detail of rosette of anisochelas-I shown on below left corner of Fig. 5b; scale equals 50 wm; d, Detail of bundle of megascleres showing abundance of sigmas around it; scale equals 50 pm. (a—d, paratype USNM 38963.) view) head 40% of total length, foot with very small alae, the frontal one ending on top in a thin digitiform process (Fig. 6d). Sigmas slender, markedly bent on hooks. Etymology.—The name citrina is derived from citrus, for the lemon-yellow color of the live sponge. Ecology.—Specimens were found under the roofs of mangrove overhangs, or, less commonly, on peat banks where they were protected from direct sunlight by neighbor- ing bushy algae (e.g., species of Halimeda, Caulerpa, Jania). Distribution.—Belize. Remarks.—This species is close to Myc- ale americana van Soest (Fig. 7), but dif- fers by the possession of a third category of anisochelas, larger megascleres and sig- mas, and lemon-yellow color. M. americana was originally described as red (van Soest 1984), but orange-yellow and olive-yellow specimens have also been reported (Zea 1987). Reexamination of Zea’s material re- vealed the occurrence of an intermediate- size category of anisochelas, overlooked by the author. This feature and the larger di- mensions of spicules in the Colombian ma- terial make us confident in assigning the Colombian specimens to M. citrina. Only one specimen quoted by van Soest (1984) in the type-series of Mycale ameri- cana was not collected in a mangrove (ZMA POR 3889, on Halimeda). Unfortu- nately the live color of the specimen was not registered but comparison with the ho- lotype (Table 3) revealed three categories of anisochelas, instead of two as originally quoted, thicker megascleres, and slightly larger sigmas. We suggest to assign this specimen to M™. citrina. Subgenus Arenochalina Lendenfeld, 1887 Diagnosis.—Mycale without ectosomal skeleton, with choanosomal skeleton of 748 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Mycale citrina, spicules: a, Terminations of subtylostyles; scale equals 10 wm; b, Face view of anisochela-I; scale equals 5 jum; c, Profile view of anisochela-II, note short and thin digitiform spur on top of frontal ala of foot (arrow); scale equals 5 jm; d, Profile view of anisochela-IIJ, note thin digitiform prolongation on top of poorly developed frontal ala of foot (arrow); scale equals 5 wm; e, Slender sigma; scale equals 10 wm. (a—e, holotype USNM 38942.) stout fibers forming a coarse rectangular re- ticulation, and with simple spicule comple- ment. Species of this subgenus are known to exude large amounts of mucus upon han- dling. Type species: Arenochalina mirabilis von Lendenfeld, 1887 (=M. mirabilis, sen- su Wiedenmayer 1989). Remarks.—The status of Arenochalina is uncertain. The diagnosis is in essence that provided by van Soest (1984) for Acama- sina de Laubenfels, 1936a. Wiedenmayer’s (1989) addition with respect to the frequent coring of fibers by foreign debris is not up- held here as this seems to be noteworthy for Australian species only. Shared traits among populations from both sides of the Atlantic, the Indo-west Pacific and southern Australia, seem to be the choanosomal, coarse, rectangular reticulation of spiculo- fibers, the low diversity of categories in the spicule complement, and the production of abundant mucus. Nonetheless, the derived condition of these characters is far from be- ing established, as suggested by similar oc- currences here and there in Mycale assigned to other subgenera. Mycale (Arenochalina) laxissima (Duchassaing & Michelotti, 1864) Figs. 8, 9, 17d; Tables 4, 5 Acamas laxissima Duchassaing & Michel- otti, 1864:95, pl. XXII, fig. 3. Esperella nuda Ridley & Dendy, 1886:339; 1887:70, pl. XV, figs. 5, 11, 14; pl. XVI, fig. 1. Hircinia cartilaginea.—Hyatt 1877:549 (not Spongia cartilaginea Esper, 1798). Hircinia cartilaginea (Esper) var. horrida Hyatt, 1877:549; pl. 17, fig. 29. Hircinia purpurea Whitfield, 1901:49, pl. 4 (not H. purpurea Hyatt, 1877:550). Mycale angulosa.—De Laubenfels 1936a: 116, fig. 2, pl. 15, 1—? Lévi 1959:129; fig. 19; pl. 6 fig. 5; Lopez & Green 1984: 749 VOLUME 111, NUMBER 4 LO-4F 1T9-VS VI cI-I1 CC-9 0C-81 CV-9 LEVE II-O6-L X 6C£E-S 60&-9LT (poinsvouwlol “688E AYOd VINZ) adAjeieg CSW AACE vI-€ €I-Cl punoj jou Ov—-6 9E-VE 8-8 9-S X 09C-6 CrC-96I (poinsvoulal “p/Or YOd VINZ) adAjojoH €9-9 Lr-€T poyiodar jou cC6 €1-Ol OV-6 9E—0E OI-cZ-S X 8EE—-9'S9C-06I (P86I IS90g UPA) DUDIAWID “~~ v-C X CL-6 49-85 e€I-Il (¢ = U) 6I-9I OV-0 9€—€E LSC X OSE-T TIE-9LT (poinsvouter) 9/70 AOd-ANI v-£ X PL 89-€9 e€I-Ol (i = 2) We IVP 8&—-CE 6-69-S X 99E-6'8CE-BIE (poinsvoulel) [770 WOd-ANI 0L-6 99-09 eIl-9 TI-Ol povodar jou LVF OF SE Co TZ-€ X ILe-l [TrE-60€ (L861 B2Z) DUDIILAUD “PW C8Z8Z-9L VIZ CclI-~Cl €c 0 1c-81 VS—C 64 SV OI-Il 8-L X VCr—-I 66&-99¢ (6€9 fUNW) L968E WNSN 98-£ 8Z-CL CIl-¢ €1-Cl SCO ECIC VSL 6F_-tV IIl-€ 6-8 X 9SV-6 CCr-80V (8€9 [ANW) 8S68€ WNSN 88-/ 08-SL VI-6 CI-CI 9C-C €C-61 CS-¥ 64 VY IIl-98-L X LLY-F LEF-OSE (LE9 [ANW) 9968 WNSN. 88-Z ZZ-69 vI-9 cll VC-F I c-0C VS-S SFI OI-1 6-8 X 19t-# 8IPF-9LE (9€9 [ANI) €968€ NNSA ; sodAjeieg 6L-6 FZ-OL vI-l €1-€l vc-9 Tc-61 8V-S SP-CE 6-9 8-8 X OSt-F C8&E—-6EE (SE9 [AUNW) Tr6se WNSN adAj0joH DUIAJII aJDIKP SPUISIS ILI se[ousosiuy II Se[oyoosiuy J sejeyoosiuy sa[Ayso[Aiqns [euajey ‘(poyeys ssojUN ‘QZ = U {SOI[e}I UT SUBS YIM “YIPIM x YISUS] IO ‘YIZUaT Jo sosueI) VUDIIIaWUD ajvIKPy pue YUIABID ajvIKP IO} (VIM UL) syUOUTOINSeoW spnoIdS—"¢ o]qQe I, 750 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. subtylostyles: scale equals 10 pm; b, Profile view of anisochela-I; scale equals 5 4m; c, Profile view of aniso- chela-III, note absence of digitiform process on top of well developed frontal ala of foot (compare with Fig. 6c); scale equals 5 4m; d, Stout sigma; scale equals 10 wm. (a—d, holotype ZMA POR 4074.) 79 (not Pandaros angulosa Duchassaing & Michelotti, 1864:89; pl. [IX fig. 4). Mycale sp.—Reiswig 1973. Thorecta horridus.—Wiedenmayer 1977: 70, pl. 8, figs. 2—4; pl. 9, fig. 1. Acamasina _laxissima.—Wiedenmayer 1977:146—147, 255. Mycale laxissima.—van Soest 1981:12.— van Soest et al. 1983:200.—Pulitzer-Fin- ali 1986:119.—Riitzler 1990:455.—Va- celet 1990:25. ?Mycale imperfecta.—Winterman-Kilian & Kahan 1984:133. Mycale (Acamasina) laxissima.—Van Soest 1984:29; pl. III, 1; fig. 9 —Zea 1987:143, WSS: Mycale mucifluens Pulitzer-Finali, 121. Mycale nuda.—Mello-Leitao et al. 1961: 12.—Hechtel 1976:254.—van Soest 1984:31.—Hajdu & Boury-Esnault 1991: 504. 1986: Spicules of Mycale americana, for comparison with Mycale citrina (Fig. 6): a, Terminations of Mycale jamaicensis Pulitzer-Finali, 1986: Sy. Mycale whitfieldi Pulitzer-Finali, 1986:127. Mycale hyatti Pulitzer-Finali, 1986:129. Material studied.—MUT POR 34, holo- type, St. Thomas, Virgin Islands. MSNG, no register number; Jamaica, Port Royal, on submerged ruins, 5—10 m, 27 Mar 1964 (M. laxissima sensu Pulitzer-Finali, 1986; num- ber PR. 40); MSNG 47695, Jamaica, Port Royal cays, 10—25 m, 22 Mar 1964 (VM. mu- cifluens Pulitzer-Finali, 1986; holotype); MSNG 47697, Jamaica, Duncans, fore-reef slope, 35 m, 30 Mar 1964 (MV. jamaicensis Pulitzer-Finali, 1986; holotype); ZMA POR 5192, Jamaica, Runaway Bay, 33.5 m, 14 Aug 1969, coll. and det. H.M. Reiswig (Mycale sp.). MCZ 7008/cat. 440, Florida, Key West; Hircinia cartilaginea sensu Hy- att, 1877 (M. hyatti Pulitzer-Finali, 1986; holotype); MCZ 7071/cat. 441, Florida, VOLUME 111, NUMBER 4 751 Fig. 8. variable dimensions (m); scale equals 500 wm; b, Detail of skeleton showing primary (1) and secondary (2) interconnecting fibers; scale equals 500 wm; c, Stout megascleres with blunt apex, and anisochelas, young megasclere is seen below center; scale equals 50 xm; d, Anisochelas of variable geometry; scale equals 20 ym. Key West; H. cartilaginea sensu Hyatt, 1877 (M. hyatti Pulitzer-Finali, 1986; para- type); MCZ 7073/cat. 222, Florida, Cape Florida; H. cartilaginea var. horrida Hyatt, 1877; M. hyatti Pulitzer-Finali, 1986; para- type. UFRJPOR 3578, Brazil, Angra dos Reis, Rio de Janairo State. USNM 41273, Belize, Twin Cays, Cuda Cut near Batfish Point, <1 m, coll. K. Riitzler, 4 Jun 1983; USNM 39281 (several lots), Belize, Twin Cays, Cuda Cut, 1 m, coll. E. Hajdu and K. Smith, 1} Aug 1993. Diagnosis.—Dark red, spiny, tubular Ar- enochalina with stout spiculo-fibers envel- oped by great amounts of spongin and forming large rectangular meshes. Choan- osome cavernous, without noticeable fleshy parts, with only one category of anisochelas and large (>50 wm) sigmas. Description.—Macroscopic appearance: Specimens are tube shaped, some in clus- ters up to six, up to 50 cm tall, 15 cm in Mycale laxissima, skeleton: a, Detail of skeleton showing fused fibers (f), and meshes of widely diameter, and thin-walled (0.5—2 cm). The color is dark wine red. A large pseudoscule (6 cm diameter) on top of the tubes is en- circled by a transparent membrane. The smaller specimens (about 6 X 6 X 6 cm) are globular and already bear an apical pseudoscule. The sponge is tough but elas- tic, compressible, spiny to the touch, and releases copious amounts of sticky mucus when squeezed. The surface is coarsely conulose from projecting spiculo-fibers. Skeleton (Fig. 8a, b): The surface mem- brane can be peeled off in places and con- tains a few scattered spicules and paucispic- ular strands of subtylostyles. The choano- somal skeleton is made up of a stout, rect- angular reticulation of spiculofibers. Meshes formed are generally within the range of 100—150 pm up to 3—4 mm (Fig. 6a). One can distinguish between primary and secondary fibers (50-300 wm thick; Fig. 8b), the latter showing clearer spongin 752 1% i 7 | - ; Fig. 9. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Mycale laxissima, spicules: a, Rare larger, and common smaller anisochelas; scale equals 10 ym; b, Common anisochela of intermediary size; scale equals 10 4m; c, Common anisochela of smaller size; scale equals 10 wm; d, Stout sigmas, and anisochelas of intermediary size; scale equals 20 pm. in transmitted light. The fibers end as sub- ectosomal, paucispicular tufts. Anisochelas are particularly abundant in a subectosomal layer of spongin where they are spread out without order. Rosettes of anisochelas are seen on the basal plate of spongin in a spec- imen from Angra dos Reis, Rio de Janeiro State, Brazil; UFRJPOR 3578. Sigmas are very common along the spiculofibers of the choanosome. Isolated megascleres and sig- mas occur in abundance, strewn at random inside the skeleton meshes. Spicules (Figs. 8c, d, 9; Table 4): Sub- tylostyles are straight, variably thick, abruptly tapering to a blunt apex, with ovoid head. Anisochelas are variable in di- mensions and geometry, generally appear narrow in profile and face views, have a curved shaft and a head 40—50% of total length. Sigmas are generally stout and Table 4.—Spicule measurements (in wm) for Mycale laxissima (fragments of different specimens) from Belize (ranges of lengths, or length < width, with means in italics; n = 10). Material Subtylostyles Anisochelas Sigmas USNM 39281 (MNRJ 640) 270—283.6—302 X 5-6.5-8 20—23.9-3 1 83—94.5-106 X 4-6 MNRJ 641 249-—272.4-292 X 5-6.1—10 22—23.6-29 83-93.3-112 X 2-3.2-6 MNRJ 642 249-277.2—-297 X 6-6.5-9 18—24.5—30 79—96.1-108 X 3-5 MNRJ 643 260—279.5—297 X 3-7.5-9 21—27.2—32 80—96.7-112 X 4-6 MNRJ 644 249-281.7—307 X 5-6.9-9 22—27.6—33 87—96.3-105 * 4—4.8-7 MNRJ 645 244—262.9-286 X 4-6.6-8 18—24.8—32 90-100.9-114 X 6 MNRJ 646 265—285.7-302 X 5-6.3-8 21—25.5—32 83—94.3-110 X 4-6 USNM 41273 265—285.1—302 * 5—5.9-7 19—25.6—34 87-97.0-112 X 4-6 23 punoj jou 9+rEYV X 901-8 88-8L 6C-8 FC-ET II-8'9-V X CCE-8 9LC VET poMseowlol payodar jou CL X SOI-SL 9C-ET CII-t x OOE-O€ET (DAY “W S¥) OR6T WeUls-sozi[Ng poyodar jou OL-IL CCC-S 81 CC X OLCVITC (Ipjayi1yM “PW Se) J “To O86] HeuLs-Joznd punoj jou €-8 CC X L8-008-9L ccs 6I-SI OVE X €8C-4 99C-6VC PoInseowior (27e1) OOT CCC X 06-$9 ec-91 C'e-€ X O8C-OST (sisuazmuvl “PF Se) 986] TWeuLq-Jozpnd [PELNOY! KOU VEC X S6-8 88-18 6c-I SCI 9-SP-£ X VOC~C 8LC-8BST pomseouwlor peyiodar jou CEC X C6-LL 6c-61 CVS e X 06C-0SC (suanpionu “W Se) O86] TeUulq-rozit[nd punoy jou 9 X CII-6 00I-9L Oc—-1 9C-CT €I-O06-S X €8C-8 6SC-SET poinseouwlol poyoda jou Oc! 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UT) syUsWOINSvaUT sfnoIdgS—¢ 2TqeL 754 abruptly bent into hooks with sharp end- ings. Ecology.—Moderately common in shad- ed marine mangrove environments with strong tidal water flow, on protected patch reefs below 5 m, and in fore-reef habitats below 12 m (compare also the detailed analysis of reef populations by Reiswig (1973) in Jamaica (as Mycale sp.). The heavy spongin fibers are known to be bored and occupied by filamentous algae (Riitzler 1990) and experimental work by one of us (KR) has shown that the coring of siliceous spicules contribute photosynthetically use- ful light levels inside the dark sponges, comparable to glass-fiber light guides in op- tical applications (similar observations for other siliceous sponges were reported by Gaino & Sara 1994, and Cattaneo-Vietti et al. 1996). Distribution.—Florida, Bahamas, Cuba, Jamaica, Puerto Rico, Virgin Islands, Neth- erlands Antilles, Colombia, Belize, Mexico, Brazil, West Africa. Remarks.—This species was subject of several misinterpretations in the literature. Esperella nuda Ridley & Dendy, 1886 from Brazil (holotype BMNH 1887.5.2.171) turns out to be a junior synonym and the originally reported (Ridley & Dendy 1887) exceptional thickness of its megascleres (16 44m) was found to be only in the range of 5—13 ym. Specimens described by Pulitzer- Finali (1986; as M. hyatti) and Zea (1987) possess comparable megascleres (4-11 wm and 2-11 wm thick; Table 5), an isolated characteristic that does not justify separat- ing a species. The species was also con- fused with Mycale angulosa (de Laubenfels 1936a, Lévi 1959), as discussed by van Soest (1984). Another erroneous identifi- cation may have been the report of M. im- perfecta Baer from Colombia (Winterman- Kilian & Kilian 1984). The suggestion of possible amphi-Atlantic distribution of M. laxissima (van Soest 1984) raises the ques- tion of apomorphies of other distant spe- cies, such as M. setosa (Keller 1889) and M. euplectellioides (Row 1911) from the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Red Sea and Indian Ocean, respectively, be- cause our examinations suggest that there is greater intra-than interspecific variation in this taxonomic complex, for instance, of an- isochelas in a single specimen. Clarification of this problem will ultimately depend on study of populations from many localities worldwide and we caution from new de- scriptions based on poorly preserved or un- representative material (e.g., related new taxa in Pulitzer-Finali 1986). The growth form of Mycale laxissima is generally tube or funnel shaped. In Belize, even in shallow water, neither encrusting nor cushion-shaped specimens were seen and early globular stages develop directly into long, thin-walled tubes. This is in con- trast to observations in Brazil (between la- tidudes 8°—24°S; EH) where encrusting forms prevail among shallow-water popu- lations of this species. The consistency of specimens varies, suggesting polymorphism in patterns of skeleton reticulation. Like- wise, abundance of microscleres varies among specimens but no attempt was made to quantify differences within and between individuals because these sponges were of- ten large and only fragments have been col- lected to allow cut specimens to regrow to protect their apparently small population Size. Subgenus Carmia Gray, 1867 Diagnosis.—Mycale without ectosomal skeleton, with loose choanosomal skeleton composed mainly of plumose, isolated spi- culofibers which are often paucispicular. Type species: Hymeniacidon macilenta Bowerbank, 1866 (=M. [C.] macilenta sen- su Topsent 1924). Remarks.—The absence of an ectosomal skeleton is likely to be a homoplastic oc- currence in Mycale because it is found in all species of Carmia and Arenochalina, and possibly also in the subgenus Mycale, (for instance, M. (M.?) lapidiformis (Ridley & Dendy), Hajdu & Desqueyroux-Fatndez 1994). As such, it is a weak character on VOLUME 111, NUMBER 4 755 Fig. 10. Mycale magnirhaphidifera, spicules: a, Set of spicules showing raphides-I (arrows mark the extrem- ities); scale equals 50 pm. b, Terminations of tylostyles, note blunt apex, and well-marked oval head; scale equals 10 pm. (a, b, USNM 39278.) which by itself to base a subgeneric diag- nosis. Mycale (Carmia) magnirhaphidifera van Soest, 1984 Figs. 10, 11, 17e; Table 6 M. (C.) magnirhaphidifera van Soest, 1984: 72 1/3 {ol UG Ve M. cecilia.—Wells & Wells, in Wells et al. 1960:212 (not M. cecilia de Laubenfels, 1936b:447). Material studied.—ZMA POR 4885, ho- lotype, Curacao. USNM 23633, North Car- olina, Hatteras Harbor, coll. and det. H. W. & M. J. Wells (as M. cecilia), 21 Nov 1959. USNM 42878, Belize, Twin Cays, Lair Channel, on red-mangrove stilt root, <1 m, coll. K. Rititzler, 10 May 1985 (dark royal- blue, sigmas rare); USNM 42979, Belize, Twin Cays, Main Channel, on red-man- grove stilt root, <1 m, coll. K. Riitzler and K. Smith, 26 Apr 1986 (purplish, with sa- bellid worm tubes; many sigmas); USNM 42949, Belize, Twin Cays, Main Channel, on red-mangrove stilt root, <1 m, coll. K. Riitzler and K. Smith, 26 Apr 1986 (dark wine red, tan below surface; no sigmas); USNM 47874, Belize, Twin Cays, Grouper Gardens entrance, on a strand of rope tied to a plastic pipe protruding from the chan- nel bottom, 1 m, coll. K. Riitzler, 1 Feb 1986 (ochre, sigmas rare); USNM 43029, Belize, Twin Cays, Sponge Haven, 1 m, coll. K. Riitzler, 26 Apr 1989 (purplish wine red; sigmas moderately common and clear- ly in 2 size classes); USNM 43040, Belize, Twin Cays, Hidden Creek, <1 m, coll. K. Riitzler, 1 May 1989 (gray, dirty yellow in- side and where dying, enveloping sabellid tubes; without sigmas); USNM 43033, Be- lize, Twin Cays, Hidden Creek, <1 m, coll. K. Riitzler, 1 May 1989 (gray; sigmas abun- dant); USNM 43036, Belize, Twin Cays, Gator Creek, <1 m, coll. K. Riitzler, 3 May 1989 (grayish ochre; sigmas very abun- dant); USNM 43037, Belize, Twin Cays, Gator Creek, <1 m, coll. K. Riitzler, 3 May 1989 (grayish ochre, massive; sigmas very abundant; with embryos); USNM 39278, Belize, Twin Cays, Hidden Creek, | m, coll. E. Hajdu & K. Smith, 9 Aug 1993 (bluish purple, sigmas common); USNM 39279, Belize, Twin Cays, Cuda Cut, | m, coll. E. Hajdu & K. Smith, 11 Aug 1993 (bluish purple, sigmas common; with larvae); USNM 39280, Belize, Twin Cays, Turtle Pond, 1 m, coll. E. Hajdu, 17 Aug 1993 (bluish purple, sigmas common); USNM 41442, Belize, Twin Cays, Cuda Cut, on red-mangrove stilt root, <1 m, coll. K. Riit- Zler, 2 May 1994 (grayish blue, very thin crust; sigmas very rare); USNM 47865, Be- lize, Twin Cays, Hidden Creek, <1 m, coll. K. Riitzler, 29 Mar 1997 (blue, sigmas abundant). Diagnosis.—Bluish-purple to brown- gray, crustose Carmia with two types of an- 756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 11. Mycale magnirhaphidifera, spicules: a, Profile view of anisochela-I, note basal portion of lateral and frontal alae of the head diverging from each other (arrows), the lateral ones projecting behind the shaft; scale equals 5 wm; b, Face view of narrow anisochela-I, and part of anisochela-II on top; scale equals 5 wm; c, Profile view of anisochela-II showing short, thin digitiform spur on top of frontal ala of its foot (arrow); scale equals 5 wm; d, Sigmas-I and -II; scale equals 10 pm. e, Raphides-I; scale equals 5 xm; f, Raphides-II; scale equals 5 wm. (a, b, d, e, USNM 39278; c, f, holotype ZMA POR 4885.) Vey, VOLUME 111, NUMBER 4 €c-E 61 V1 61-7 9I-E1 Ics 8I-S1 oI-T FI-11 Le-€ 6I-V1 8C-6 OC-EI 6I-OZI-SI CCS LI-El (p = 4) OC-II (LZ = 4%) 07-891 poevodar jou Il 46-8 II seprydey O8c—l €I1c-101 98C-8 FI C-6Cl SSc-9 C6I—-601 CSC-E FS I-S6 O9CZ ESI—-0CI 6vc-8 681-11 CCL—6 VOE-08C vcr F106 poevodeai jou Ole—€ 16c—097C | soprydey GC] ce punoj jou (1 = 4%) 07 (1 = 4%) 6 9C-0'€c-61 punoj jou punoj jou (€ = Y) IC-€I punoj jou poviodoar jou punoj jou [J SPUISIS 8v-6rr-LE etI-Zcl-Cl 9V-C EF-OV CI-OCI-II Lv-OSt-Cy 8I-67I-I1 8r-l €r-Ov 9I-T rI-Il Lv-LE&r-Le €l-cll-ol CS-I Lr-Sv eI-€TI-6 (= 4) I€9C SISVSI-CI OS-9 EF-CE eI-I1 SS-l6r-9b 8I-9'SI-EI 8S—0S oI-+1 REO OC LI-¥cl-Ol ] SeUISIS II sejeyoosiuy ve-8 TE-TE 9-8 CE-IE (¢ = 4) LE-€E NE GENE ve—-9 TE—-6T (¢ = 4) 9€-6¢ Ser rc-ITC (y = 4) SE-lE Se-6 OF VE 9e-0€ ee -9 Lc--61 ] sejeyoosiuy T9-Cv X POC-O0'SLC-8ET VS-CV X OOE—C 6LZC-SST 99-V X CCL—-O'EDE-VLT €S-VV X €8C-F 69C-CET CS-6E X 80E-1 E8C-8ST OS-6'€ X 80E—-O0'8ZLC-CET O9-VV X V6C-9 99C-CET VV X 98C-C 8S c-961 9-v X 8SC-O0 TEC 961 9-V X OLC-OSTC €-CC-S I X OLC-C HS C-9ET sa[Ayso[Aiqns EEE vS9 faNW (€S9 [AUNW) O876E WNSN cS9 CLANW 1S9 fUNW OS9 LANA 679 CANW 879 CANW (Lv9 TANI) 8Z76€ WNSN azi[og poinsvoulol (011999 “W S® “€€9ET IWNSN) O96 SITSM 7 SITS eulforeD YON (S88r HOd VINZ) 2dA0[0H [eae ovseing ee Dene, EEE ‘(payeis ssayun ‘QZ = u ‘SOI[eII UI SURO YIM ‘IIPIM x YSU] JO ‘yISue] Jo sasuv1) Diafipiydvysusou avd 10F (uM ul) syuouonsvow! spnoidg—9 2IqGeL 758 isochelas, with or without sigmas, and with two size classes of raphides, the larger ones reaching more than 200 wm in length. Description.—Macroscopic appearance: Encrusting, up to (5 mm thick) very soft and fragile. Subectosomal channels of the aquiferous system converging to oscula (S3 mm diameter) which are raised and sur- rounded by transparent collars. Color in life is highly variable (see material section above), burgundy to bluish-purple and grayish blue in light-exposed areas, cream to yellowish in shaded zones; drab in dying tissue parts. Most specimens collected (May to August) contained larvae (150-300 wm in diameter) in the choanosome; many were associated with clusters of serpulid poly- chaete tubes. Skeleton: The ectosome, lacking a spe- cial skeleton, consists of a superficial mem- brane bearing pores (30-50 pm diameter) and is supported by terminal brushes of choanosomal spiculofibers. A few sigmas and single anisochelas are seen between the pores, rosettes of anisochelas I (ca. 70 wm diameter) occur in the subectosomal zone. The choanosome has a low spicule den- sity, with subtylostyles organized in pauci- spicular (3 to 10) ascending tracts of 15-30 1m in diameter. The fibers branch seldomly and generally run parallel to each other without anastomosing. Close to the surface they diverge to form brushes. Sigmas are very common throughout the sponge. Raphides too are dispersed, not organized into trichodragmas. Spicules (Figs. 10, 11; Table 6): Subty- lostyles straight, slender, blunt, with char- acteristic irregularly-oval head. Anisochelas I, narrow in face and side views, head about 50% of total length, base of frontal ala of the head curved forward, base of lateral alae of the head curved backward and slightly protruding over the shaft (in side view). Anisochelas II slender, head about 50% of total length, frontal ala of foot ex- tended on top in a short digitiform process. Sigmas I with almost straight inner face, abruptly bent into very sharp hooks. Sigmas PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON II delicate with sharp points. Raphides I slender, straight, sharply pointed. Raphides II stout, straight, sharply pointed. Ecology.—A common species in the sur- vey area where it prefers shaded substrates. Many specimens were found on the bare tips of new Rhizophora mangle prop roots suggesting capacity to quickly colonize newly available substrate (R-strategist). The common association with polychaete worms was also observed by Wells etal. (1960, as M. cecilia) and van Soest (1984). Distribution.—North Carolina, Curagao, Belize. Remarks.—In contrast to the holotype from Curacao, most Belizean specimens contain abundant sigmas (see material sec- tion above). On the other hand, color, shape, and morphology of all spicules other than sigmas agree well, including the sec- ond smaller category of raphides that was found in the holotype (Fig. 11f) but was not originally reported (van Soest 1984). Nev- ertheless, the abundance of sigmas in some specimens is striking (see material section above) and may justify separating the sig- ma-bearing population as a different species if more supporting data become available. The material from North Carolina iden- tified by Wells & Wells, in Wells et al. (1960) as Mycale cecilia has previously been referred to both M. microsigmatosa (Hechtel 1965) and M. americana (van Soest 1984). We have analyzed the speci- men (USNM 23633) and found it conspe- cific with M. (C.) magnirhaphidifera, also bearing sigmas (Table 6). Wells & Wells, in Wells et al. (1960) described their material as pale yellowish-green or yellowish-tan, similar to the color of M. (C.) magnirha- phidifera from shaded habitats. Mycale (Carmia) microsigmatosa Arndt, Sn 7) Rigs. 12) 135 17> Table 7 Mycale fistulata var. microsigmatosa Arndt, 1927:144, pl. I, 9. Mycale_ microsigmatosa.—Burton 1956: VOLUME 111, NUMBER 4 I Fig. 12. 759 Mycale microsigmatosa, spicules: a, Terminations of subtylostyles showing heads, and gradually sharpening apex; scale equals 10 wm; b, Terminations of subtylostyles showing head and gradually sharpening apex; scale equals 10 pm. (a, USNM 33580; b, holotype ZMA POR 1593.) 129.—Hechtel 1965:47.—van Soest 1981:12.—Winterman-Kilian & Kilian 1984:132.—Pulitzer-Finali 1986:124.— Riitzler 1986:120. Mycale (Carmia) microsigmatosa.—van Soest 1984:24; pl.II, fig. 6—Zea 1987: 142. Not sensu Green & Gomez 1986: 284 (=M. cecilia de Laubenfels). Carmia microsigmatosa.—Hajdu & Boury- Esnault 1991:510. Material studied.—ZMA POR 1593, ho- lotype, Curacao. USNM 22207, holotype of M. cecilia, Panama City, Panama. USNM 33580, Belize, Twin Cays, Turtle Pond en- trance, on root of Rhizophora, coll. I. Goodbody, 24 Feb 1985; USNM 34625, Belize, Twin Cays, coll. I. Goodbody, Feb 1986; USNM 41258, Belize, Twin Cays, Main Channel south of dock, on mangrove roots, coll. K. Riitzler, 12 Jun 1983; USNM 39302 (several lots), Belize, Twin Cays, Cuda Cut, coll. E. Hajdu & K. Smith, 13 Aug 1993; USNM 39326 (several lots), Be- lize, Twin Cays, Sponge Haven, coll. E. Hajdu & K. Smith, 9 Aug 1993; USNM 47873, Belize, Blueground Range, <1 m, coll. K. Riitzler, 6 May 1994. USNM 42951, Belize, Twin Cays, Batfish Point, <1 m, coll. K. Riitzler, 19 Apr 1996. Diagnosis.—Orange-dotted to reddish, crustose Mycale (Carmia), with two cate- gories of small (<30 wm) anisochelas and small (<50 wm) sigmas. Description.—Macroscopic appearance: Specimens are encrusting or cushion- shaped (5 cm thick), very soft and fragile. Subectosomal channels of the aquiferous system converge to scattered oscula (<5 mm) with transparent, membranous collar. Live color is highly variable but predomi- nantly reddish orange. Even grayish-green- ish specimens have recognizable bright or- ange specks which aid in field recognition of the species. Specimens with larvae were found during August. Skeleton: A thin surface membrane with scattered megascleres and a few paucispi- cular (3 to 6 spicules wide) bundles can be peeled off and terminal spicule brushes of the ascending choanosomal fibers reach and penetrate the ectosome. The choanosome is supported by mean- dering tracts of subtylostyles (50 wm thick) which may be more or less replaced by for- eign structures, such as algae and sabellid polychaete tubes. These tracts split up and anastomose near the surface where they fan out in brushes. Scattered megascleres are common throughout and sigmas are seen here and there in large concentrations. Spicules (Figs. 12, 13; Table 7). Subty- lostyles slender, mostly straight, with faint- ly marked oval head, apex sharpening grad- ually or abruptly. Anisochelas I, head about 60% of total length, shaft gradually curved, basal portion of frontal and lateral alae PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 13. Mycale microsigmatosa, spicules: a, Profile view of anisochela-I; scale equals 5 jm; b, Profile view of anisochela-II; scale equals 5 ym; c, Profile view of anisochela-II showing thin digitiform process on top of frontal ala of foot (arrow); scale equals 5 pm; d, Slender sigma; scale equals 10 um; e, Profile view of anisochela- I; scale equals 5 xm; f, Profile view of anisochela-II showing thin digitiform process on top of frontal ala of foot (arrow); scale equals 5 ym; g, Stout sigma; scale equals 10 wm. (a, b, d, USNM 33580; c, holotype ZMA POR 1593; e—g, holotype of M. cecilia, USNM 22207.) 761 VOLUME 111, NUMBER 4 6£-S SE-CE CI-I FI-el 0c-06!-81 V-€ X 69C~C 6FC-ICC L99 CANIN SEL LESCE 91-9 FI-El €c00c-81 9-S F-E X VLC-F SSC-VCT 999 fTANW 6£-6 SE-IE (9 = 4) rI-TI Ic-c 0c-81 L-O'S-V X 6SC~C OFC -OET $99 fANW ev-0LE-CE CI-8 €1-El Ics 61-81 9-V X 88C—-1 99C-EVT (y99 fUNW) 97E6€ WNSN 6£ fF FE-IE (C= #9) CI Oc-9 8I-LI 9-C re X LOC-LISC-TCET £99 fANW 6€-8 LE-9E Cl-¥ €I-€l Occ 8I-LI 9-Cr-E X BSCS OFC-LTETC c99 TANW 6£-Z SE-CE Cl-- €I-Tl 0c-8 81-81 c-€ X €LC-O'CSC-8ET 199 fANW Cy -99E-TE CIl-8 rIV1 €C-9 0C-81 9-V X 88C-L L9C-9VT 099 fAaNW 6£-8 FE-CE CI-8 €1-el Oc Z8I-81 9-V X £9C-OLPC-SET 689 fANW Iv-6 9E-CE SI-8 €1~Cl VC-E 0C-61 L-LE-V X 88C-9 69C VVC 859 CANIN CE-C €E0E 9I-€ FI V1 1c-96I-LI 9-lr-€ X LLC-L6EC-YITC LS9 fANW LE-O'SEVE SI-s €1-Cl 1c-6 SI-LI C-I CC X 60C-6 YEC VCC 999 CANIN LE-8 EE-6C (€ = 4) vI-€l Oc-S 61-81 S-€ X VLT-8 ESC-SET (SS9 fUNW) ZOE6E WNSN Cy-&£ 8&-0E (= 2) II 0c-98!-81 p-€ X 09T-9'I#Z—-917 8Sclv WNSN 9€-8 EEST SIO +rI-Cl 0c—06!I-81 L-9 X CSC PH LEC-EIT Score WNSf) Cy-0 SE-OE CI-l rI-€l Ic-€ 6I-81 9 X £9C-6 IPC-VCET O8sce NNSA Le’ CE-OE vI-€ €I-El 0c-0 81-91 9-V X €8C-8 09C-TET poinseoulol 8E—CE UOATS JOU 61-8 8 X v6c—OSTC (L761 IPUIV “E6ST WOd WWNZ) 2dM0[0H SeUISIS II se[eysosiuy ] sejeyoosiuy satAysopAiqns [earl ee ee ee ee ‘(pajejs ssoyun ‘QZ = u {soI[eII UT SUBOU WIM ‘“YIPIM x YISUI] 10 ‘YIZU] Jo sosuLI) VSOJDUISIsOsONU a]DIKP IOJ (WIA UT) sJUoWIOIMSeoUN s[NoIdg—Y/ IIqQRL 762 slightly divergent. Anisochelas II, head about 50% of total length, shaft gradually curved, with basal portion of frontal and lateral alae of head slightly divergent; in some of these spicules the top of the frontal ala of the foot tapers to a thin digitiform process (Fig. 13f). Sigmas slender, both ends markedly bent into sharp hooks. Ecology.—Common on mangrove prop roots, including new growth tips, and all other light-exposed substrates in shallow la- goon and harbor habitats (R-strategist). The species is also reported as tolerant of oil and domestic wastes (Muricy 1989). Sabellid polychaetes are commonly associated with and overgrown by this sponge. Distribution Bermuda, Florida, Baha- mas (?), Cuba (?), Jamaica, Puerto Rico, Venezuela, Netherlands Antilles, Colombia, Belize; also from Brazil, West Africa (7?), Acores (?), and Eastern Mediterranean (7). Remarks.—Agreement with the holotype (Figs. 12b, 13c) is excellent, including the presence of two categories of anisochelas which has not before been reported for the species. Because anisochelas are rare in this species (van Soest 1984, Hajdu & Boury- Esnault 1991), it is not surprising that oc- currence of a second category was previ- ously overlooked. Mycale microsigmatosa is very common in Belizean mangroves, displaying mainly reddish-orange color but also red and trans- lucent gray. Similar color variability was reported by Hechtel (1965, Jamaica), van Soest (1984; Curagao, Bonaire, Margarita, and Florida), and Zea (1987, Colombia). Hajdu & Boury-Esnault (1991) found Bra- zilian specimens (collected between lati- tudes 13° and 24°S) to be exclusively red but subsequent observations (EH) found pale-yellow specimens from Angra dos Reis (Rio de Janeiro state). Bergquist (1965) and Hechtel (1965), suggested the synonymy of Mycale micro- sigmatosa with M. cecilia de Laubenfels. Another opinion by van Soest (1984) fa- vored a close relationship between M. ce- cilia and M. americana. The present finding PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of two categories of anisochelas in M. mi- crosigmatosa rather supports the former view, agreement with M. cecilia (holotype, Fig. 13e—g). Furthermore, de Laubenfels (1936b) describes M. cecilia as “‘basically green, abundantly provided with bright red specks”, similar to our own observations on M. microsigmatosa. However, we prefer to keep M. cecilia as a valid name for east- erm Pacific records of the sponge, basing this decision on geographic separation and small differences in spicule dimensions. Therefore we propose to assign the record of M. (C.) microsigmatosa from the Pacific coast of Mexico (Green & Gémez 1986) to M. cecilia, this sponge has relatively thick subtylostyles (5-9 wm) and anisochelas with a large range of length (12—21 wm), possibly representing two categories. Van Soest (1984) also suggested possible syn- onymy of Mycale microsigmatosa with M. senegalensis Lévi, 1952, M. sanguinea Tsurnamal, 1969, and Desmacella melior- ata Wiedenmayer, 1977. To this list we may add Desmacella janiae Verril, 1873, Biem- na microstyla de Laubenfels, 1950a, and Prosuberites scarlatum Alcolado, 1984, but formal decision must await re-examination of all types. Subgenus Mycale Gray, 1867 Diagnosis.—Mycale with ectosomal skeleton of confused tangential megascleres over perpendicular, dendritic choanosomal spicule tracts that end in subectosomal brushes. Type species: Hymeniacidon lin- gua Bowerbank, 1866 (Hajdu & Desquey- roux-Fatindez 1994). Remarks.—A subgenus Anomomycale was erected by Topsent (1924) for Desma- cidon titubans Schmidt (1870). This sponge, however, has most characteristics of the subgenus Mycale, including the su- bectosomal brushes, large megascleres (one size class >700 ym), and possibly tangen- tial subtylostyles in the ectosome (Lund- beck 1905). Only the distinctive shape of its anisochelas separates it. This single fea- VOLUME 111, NUMBER 4 Fig. 14. Mycale laevis, skeleton: a, Transverse section showing terminally divergent spicule brushes (s) of ascending choanosomal tracts supporting the tangential ectosomal skeleton (e); scale equals 200 ym; b, Detail of surface showing tangentially disposed megascleres, and abundance of microscleres; scale equals 100 um; c, Detail of subectosomal area showing typical patches of trichodragmas, and also of isolated raphides; scale equals 50 wm; d, Fusiform, bent subtylostyle, note abundance of trichodragmas; scale equals 100 ym. ture is insufficient to justify subgeneric sep- aration and it seems appropriate to keep M. titubans in the subgenus Mycale. Mycale (Mycale) laevis (Carter, 1882) Figs. 14, 15, 17g; Table 8 Synonymy, see Hajdu & Desqueyroux- Fatindez (1994); in addition: Mycale fusca.—Mello-Leitao et al. 1961: 12; Hechtel 1976:254; Hajdu & Boury Esnault 1991:504. Mycale (Mycale) laevis.—Lehnert 1993:54. Material studied.—USNM 41267, Be- lize, Twin Cays, Sponge Haven, <1 m, on mangrove roots, coll K. Riitzler, 13 June 1983; USNM 39328 (3 lots), Belize, Blue Ground Range, | m, coll. E. Hajdu & K. Smith, 12 Aug 1993. Diagnosis.—Orange to yellow, encrust- ing or massive Mycale with large (>500 14m) megascleres and two size classes of anisochelas; with basal, spur-like projec- tions on the smaller anisochelas; also pres- ent, two classes of sigmas and raphides in trichodragmas. Description.—Macroscopic appearance: Thick cushions (up to 5 cm) or clusters of 20 or more branches (1—10 cm diameter, up to 50 cm tall) rising from a massive base and often with apical oscula (up to 4 cm diameter). Oscula have fibrous-looking col- lars made up of tracts of megascleres. Con- sistency is rough but compressible and the texture rough. Color in life is a bright or- ange-yellow, but also a few whitish speci- mens were seen. Specimens with embryos Were seen in August. Skeleton (Fig. 14a—c): The ectosome is reinforced by a thin layer of tangentially strewn subtylostyles interspersed with abundant microscleres. A very dense, con- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 15. Mycale laevis, spicules: a, Face view of anisochela-I; scale equals 20 um; b, Profile view of anisochela-I, note slight s-shape of shaft; scale equals 20 um; c, Detail of foot of anisochela-I shown in Fig. 15b, with an undulated basal portion of the falx; scale equals 5 wm; d, Face view of rare, robust anisochela-III showing thin digitiform prolongation on top of frontal ala of foot (arrow up), and basal spur (arrow down); scale equals 5 um; e, Profile view of typical anisochela-III showing basal spur (arrow), note reduced foot without alae; scale equals 5 xm; f, Sigmas-I and -II; scale equals 10 wm; g, Trichodragmas-I and -II; scale equals 20 > zm; h, Rare trichodragmas-II; scale equals 5 wm. VOLUME 111, NUMBER 4 Spicule measurements (in pm) for Mycale laevis (ranges of length, or length < width, with means in italics; n = 20, unless stated). Table 8. Sigmas I Sigmas II Trichodragmas I Trichodragmas II Anisochelas II Anisochelas I Subtylostyles Material 15—20.2—23 74—-77.4-86 not found 29-42.0-50 X 2-3 18-18. 7-21 78-83.5—89 429-504.6-551 USNM 39238 (MNRJ 668) x 8-12 72-80. 1—87 x 6—-15.9-20 509—550.7—-610 not found 31-39.5-47 X 2 18—22 (n = 5) 19-22. 1-33 72-81.8-95 MNRJ 669 x 4-8.5-17 x 9-16.0-23 525-566.0-588 x 13-15.8—21 514-560. 7-610 xX 12-14.2-18 72-79.6-90 11—/3.0-15 14—16.5—21 17—24. 7-33 32-39-49 X 3 72-80. 1-89 MNRJ 670 xX 4-5.9-8 13—17.6—26 x 8-10 70—78.4-85 17-20. 1—24 17—15.5—28 32-43.3-52 X 3 73—83.4-91 USNM 41267 x 3-4.8-8 xX 5-9.2-15 765 fused subectosomal reticulation of multi- spicular subtylostyle tracts (up to 300 um thick) is oriented mainly parallel to the sur- face. Spaces between tracts are filled by large amounts of scattered subtylostyles, anisochelas I, and patches of trichodragmas I (by far the most common microscleres, Fig. 14c). Brushes of subtylostyles support the ectosomal network (Fig. 14a), and pierce the surface slightly. Anisochelas I form a few rosettes (160 wm) but more of- ten occur scattered or surrounding the bun- dles of megascleres. The choanosome shows high spicule density and includes criss-crossing subty- lostyle tracts with loose subtylostyles and patches of numerous trichodragmas I in be- tween. Digitiform processes of the larger Specimens are supported by longitudinal spicule tracts and a secondary, transverse plumo-reticulate skeleton. Spicules (Figs. 14d, 15; Table 8): Sub- tylostyles fusiform, mostly bent with faintly marked, oval head and abruptly sharpened, or strongyloid apex. Anisochelas type I, with shaft slightly s-shaped, head ca. 40% of total length, basal portion of falx on foot undulated (Fig. 15c). Anisochelas type III slender, head 70% of total length, foot often reduced (lacking alae) and with basal spur. Sigmas I stout, with sharply bent hooks Sigmas II similar to I but more slender. Tri- chodragmas I; trichodragmas II. Ecology.—This is a well known reefal species (Goreau & Hartman 1966) and is widely distributed in the shallow parts (16— 25 m) of the fore-reed of Carrie Bow Cay (see Riitzler & Macintyre 1982 for the bion- omy of the reef). In the mangroves, Mycale laevis is restricted to areas with good ex- posure to the open, near-reef lagoon (it is common at Blue Ground Range and Cat Cay Lagoon, but very rare at Twin Cays). There, however, the sponges attached to red mangrove roots can grow to record size. Specimens laying unattached on the bottom (presumably fallen from the roots) and in- tergrown with turtle grass survive well and appear healthy. 766 Distribution.—Cuba, Jamaica, Domini- can Republic, Puerto Rico, Venezuela, Co- lombia, Belize, Mexico; also Brazil. Remarks.—A few details should be add- ed to former descriptions of the species. Notably, we recognize two categories of sigmas (smaller one can be rare) rather than a large size range of one, and two catego- ries of trichodragmas in some of the spec- imens (again, the smaller being rare see Hajdu & Desqueyroux-Fatindez 1994, tab. 1, and references therein). No taxonomic importance is given to these characters here. Esperia massa Schmidt, 1862 sensu Schmidt 1870:58 and Mycale massa (Schmidt, 1862) sensu Winterman-Kilian 1984:133, are likely to be synonyms of Mycale laevis because the latter species was originally described from the Mediterra- nean Sea and a West-Indian distribution is highly unlikely despite the similarities in skeleton morphology. Subgenus Paresperella Dendy, 1905 Diagnosis.—Mycale with an ectosomal skeleton of reticulated tangential mega- scleres and with serrated sigmas among the microscleres. Type species: Esperia serra- tohamata Carter, 1880. Remarks.—Van Soest (1984), based on the very peculiar characteristics of serrated sigmas, considers Paresperella a monophy- letic group. Nevertheless, in the absence of another congruent diagnostic character, Pa- resperella may fall to Aegogropila (see comments on Anomomycale, above, under subgenus Mycale). However, until further study of species in Paresperella and other closely related subgenera (Aegogropila, Carmia), it is convenient to maintain this subgenus for the 20 or so species already described with serrated sigmas. Mycale (Paresperella) species indet. Fig. 16 Material studied.—Spicules found con- taminating sample of Mycale_ citrina, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON USNM 38967, Belize, Twin Cays, Cuda Cut, 1 m, coll. E. Hajdu & K. Smith 13 Aug 199). Spicules.—Sigmas (Fig. 16), slender, o- or s-shaped, shaft subcylindrical (flattened in cross section), with ca. 10 spines each along the outer bends of the hooks; spines single or double and directed toward spic- ule center. Remarks.—These distinctive spicules are the only material found, thus making spe- cies diagnosis impossible. The sigmas came in abundance from a very small specimen or fragment (used up in making the prepa- ration) inadvertently collected as part of a specimen of M. citrina. Comparison with the only Paresperella species described from the western Atlantic, Mycale (P.) spinosigma (Boury-Esnault 1973; MNHN D NBE 968), indicates that the latter is most likely contaminated with Paresperella spicules. The anisochelas re- ported by Boury-Esnault (1973) were not found in a preparation of the type specimen and sigmas were very rare. The ZMA col- lection has a very small macerated speci- men (ZMA 5389) from inside Cura¢ao (Barbara Beach, 1—3 m) which, from the appearance of serrated sigmas may be con- specific with the Belizean material. The Cu- ragao specimen has slender subtylostyles (270-323 wm long), one category of ani- sochelas (24—30 ym long), and the serrated sigmas (78—98 wm long). One toxa (34 pm long) was also seen but may be foreign. We refrain from making an identification until more material is available. Discussion Some names associated with Caribbean Mycale are still in need of a reappraisal. Winterman-Kilian & Kilian (1984) listed M. cleistochela flagellifer Vacelet & Vas- seur, 1971 (a species from Madagascar) in Colombia, with sigmas twice the size of the original report. We have not examined the Colombian material but assume that it is conspecific with M. (Ae.) diversisigmata VOLUME 111, NUMBER 4 767 Fig. 16. Mycale (Paresperella) sp., characteristic sigmas (other spicules belong to M. citrina): a, Serrated sigma; scale equals 20 wm; b, Serrated sigma showing non-cylindrical shaft, note narrower middle portion when compared to wider hooks; scale equals 20 wm; c, Detail showing spines on hooks of serrated sigmas, note heterogeneous distribution with the occurrence of twins (arrows); scale equals 5 wm. van Soest, 1984 (from Curagao), which contains flagelliform sigmas (150—200 zm), aS well as semi-closed anisochelas. Similarity of the latter species with M. cleistochela flagellifer was suspected by van Soest (1984) who must have been un- aware of the morphology of true cleisto- cheles which was only recently revealed by scanning electron microscopy (Pulitzer-Fin- ali 1996:fig. 15; for M. (Ae.) peculiaris). Schmidt (1870) cited five species from Florida under genera Esperia Nardo, 1833 and Desmacidon Bowerbank, 1864, that were referred to Mycale by subsequent au- thors. The specimen originally named Es- peria renieroides Schmidt (1870) was re- ferred to Oxymycale by van Soest (1984) and to Mycale by Pulitzer-Finali (1986). Neither author had examined Schmidt’s preparation (BMNH 1870.5.3.31) which re- veals an isotropic reticulation of small ox- eas, part of a haplosclerid sponge, and a single rosette of anisochelas of the type that is indicative of M. immitis (Schmidt 1870). The specimen identified as Esperia massa Schmidt, 1862 (sensu Schmidt 1870) is considered dubious by Topsent (1924), and unrecognizable by van Soest (1984). Win- terman-Kilian & Kilian (1984) report M. massa from Colombia. However, data in- dicate that these specimens are conspecific with M. laevis which has indeed very sim- ilar spiculation. As argued above, Desmacidon titubans described by Schmidt (1870) is best iden- tified as Mycale (M.) titubans, although the conspecificity of populations from Florida (Schmidt 1870), the northeastern-Atlantic (Lundbeck 1905, Topsent 1924) and the Antarctic (Boury-Esnault & Van Beveren 1982) is certainly unlikely and needs to be confirmed. Esperia diaphana Schmidt (1870; material examined: dissociated spic- ules mount made by Schmidt, BMNH 1870.5.3.32) and Esperia immitis Schmidt (1870; material examined: dissociated spic- ules mount made by Schmidt, BMNH 1870.5.3.30, MCZ No. 42; label could be in Schmidt’s handwriting but depth of 274 m differs from that of 230 m in the original 768 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 17. Underwater photographs of live Mycale specimens representing seven of the eight species found at Twin Cays: a, M. arndti, grayish-colored specimen, X1.0; b, M. carmigropila new species, on red-mangrove stilt root, 0.9; c, M. citrina new species, on mangrove peat, X1.3; d, M. laxissima, on red-mangrove stilt roots, x0.3; e, M. magnirhaphidifera, overgrowing serpulid polychaete tubes on red-mangrove stilt roots, 0.5; f, M. microsigmatosa, 1.0; g, M. laevis on red-mangrove stilt root, <0.3. VOLUME 111, NUMBER 4 account) are regarded as good species of Mycale (Mycale). Esperia diaphana has two categories of megascleres (721-827 zm and 509—604 pm), three categories of anisochelas (148-170 wm, 31—40 pm, 19— 25 pm), one category of sigmas (16—23 um), and one category of trichodragmas (41-70 wm), and seems close to M. (M.) grandis Gray, 1867 (Indo-west Pacific), and M. (M.) anisochela Lévi, 1963 (south and southwestern Africa). Abundant material from the Gulf of Mexico, agreeing with M. (M.) diaphana is in the USNM collection. M. (M.) immitis has two categories of me- gascleres (996-1198 p.m and 466—530 pm), three categories of anisochelas (90—105 pm, 29-33 pm, 20—25 pm), one category of sigmas (35—45 wm), and two categories of trichodragmas (90-108 wm, 18-33 pm). Van Soest (1984) considers Little’s (1963) Mycale macilenta from the Gulf of Mexico to be a likely new species in the subgenus Aegogropila. In the account be- low we will treat it as Mycale (Ae.) species. The following key to 17 recognized spe- cies of Mycale of the Caribbean should be used in conjunction with the relevant liter- ature (most recent reliable or available ref- erence). Mycale (Aegropila) van Soest 1984 americana van Soest M. (Ae.) angulosa (Duch. & Mich.) M. (Ae.) arndti van Soest van Soest 1984 van Soest 1984 M. (Ae.) carmigropila this paper new species M. (Ae.) citrina this paper new species M. (Ae.) diversigmata van Soest 1984 van Soest M. (Ae.) escarlatei Haj- Hajdu et al. 1995 du et al. M. (Ae.) species indet. Little 1963 (as M. macilenta) M. (Arenochalina) lax- this paper issima (Duch. & Mich.) M. (Carmia) magnirha- Phidifera van Soest van Soest 1984 769 M. (C.) microsigmatosa Arndt M. (Grapelia) ungui- fera Hajdu et al. M. (Mycale) diaphana (Schmidt) this paper Hajdu et al. 1995 Schmidt 1970 (as Esperia); this pa- per (discussion) Schmidt 1970 (as Esperia); this pa- per (discussion) this paper Schmidt, 1870 (as Desmacidon); this paper (comment, subgenus Mycale) this paper M. (M.) immitis (Schmidt) M. (M.) laevis (Carter) M. (M.) titubans (Schmidt) M. (Paresperella) spe- cies indet. Key to the species of Caribbean Mycale 1. Megascleres near 500 pm common .. 2 Megascleres always <500 pm ...... 5 2. Two categories of megascleres ...... 3 One category of megascleres, trichod- ragmas often abundant Mek ating ila, eee hc Tecan Mycale (M.) laevis 3. Three categories of anisochelas, aniso- chelas I >60 pm One category of anisochelas <60 pm, twisted M. (M.) titubans 4. Anisochelas I =150 pm common .... Be Ge Sika 2 oases Ee LENS Pere M. (M.) diaphana Anisochelas I <110 wm, with shaft markedly curved in profile view MARE Cee nae cae esos M. (M.) immitis Sa Sethated sigmas Present sa. sae Sls ute M. (Paresperella) species indet. Serrated sigmas absent ............ 6 6. Three or more categories of anisochelas 7 One or two categories of anisochelas ... 12 Pe Sieimasspresentgasa. 1.29 oebaces ots ee 8 Sigmas absent, two categories of toxas M. (Aegropila) species indet. 8. Two or more categories of sigmas ... 9 One category of sigmas............ 10 9. Two categories of sigmas, anisochelas II very narrow (“‘duck’s-bill’’) dotnet duly ian hit aE eau oe at Aa M. (Ae.) escarlatei Two categories of flagelliform sigmas, three of normal ones BERRY cr GUA eens M. (Ae.) diversisigmata Three categories of sigmas, four cate- gories of anisochelas .... M. (Ae.) arndti 10. Anisochelas only palmate 770 Anisochelas I and II unguiferate, ani- sochelas III palmate with spur Uelee Sat CaS Be ie M. (Grapelia.) unguifera 11. Sigmas 270 wm common, live sponge lemon-yellow M. (Ae.) citrina Sigmas <70 pm, live sponge blue AVEO DW ANA ey tases M. (Ae.) carmigropila 1D aRaphide sic omin@n eae ee ere ee 13 Raphides absent or very uncommon (possibly contaminants) ............ 14 13. Two categories of raphides, raphides I >100 pm, isochelas absent Be peareg Mere M. (Carmia) magnirhaphidifera One category of raphides <50 pm, is- ochelas present ....... M. (Ae.) angulosa 14. With tangential ectosomal reticulation Er STO ey eee M. (Ae.) americana Without tangential ectosomal skeleton 15 15. Sigmas common, >50 wm, coarse choanosomal spongin fibers, massive sponges .... M. (Arenochalina) laxissima Sigmas <40 pm, delicate choanosomal skeleton strands, encrusting sponges . . sae seme es. ae Se M. (C.) microsigmatosa Acknowledgments We are indebted to EF R. Schram and R. W. M. van Soest (Amsterdam) for critical review of early drafts of the manuscript. L. B. Holthuis (NNM) clarified doubts con- cerning the type species of Aegogropila. A. B. Johnston (MCZ), C. Lévi (MNHN), V. Rainieri (MSNG), S. Stone and C. Valen- tine (BMNH), and S. Zea (INV) are thanked for the loan of specimens. C. Val- entine is specially thanked for her efforts on behalf of Johnston’s Halichondria ae- gagropila. K. Smith (National Museum of Natural History, Washington, D.C.) assisted in field work in Belize and in laboratory preparations; she is credited for finding a new species, Mycale (Aegogropila?) car- migropila. We thank M. K. Ryan (National Museum of Natural History, Washington, D.C.) for preparing Fig. 1 and helping with other illustration chores, and C. C. Hansen (also from the Museum) for taking and printing the photograph in Fig. 17d. E Hiemstra (Amsterdam) is thanked for ad- vice on SEM preparation. EH was support- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ed by fellowships and grants from CNPq and FAPESP, Brazil. His field work in Be- lize was made possible by a short-term vis- itor grant from the Smithsonian Office of Fellowships and Grants and by a fellowship from the Smithsonian’s Caribbean Coral Reef Ecosystems program (CCRE; partly supported by a grant from the EXXON Corporation), National Museum of Natural History. This is CCRE contribution number Sole Literature Cited Alcolado, P. 1984. Nuevas especies de esponjas en- contradas en Cuba.—Poeyana 271:1—22. Arndt, W. 1927. Kalk- und Kieselschwamme von Cu- ragao.—Bijdragen tot de Dierkunde 25:133- 158, pls. I-HI. Bergquist, P. R. 1965. The sponges of Micronesia, part 1. The Palau Archipelago.—Pacific Science 19: 123-204. , & J. P Fromont. 1988. The marine fauna of New Zealand: Porifera, Demospongiae. Part 4 (Poecilosclerida).—New Zealand Oceanograph- ic Institute Memoir 96:1—197. Boury-Esnault, N. 1973. 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First record and new species of Gastrophanella (Porifera: Demospongiae: Lithistida) from the central East Pacific Patricia GOmez Instituto de Ciencias del Mar y Limnologia, U.N.A.M. Circuito Exterior S/N, Ciudad Universitaria, C.P. 04510, México, D.F Abstract.—A new species of sponge, Gastrophanella primore (Lithistida, De- mospongiae), is described from the continental shelf off Guerrero, Mexico, cen- tral East Pacific at a depth of 45 m. This is the first record of a species of Gastrophanella and the second member of the Lithistida reported from the East Pacific. Its cup shape, tight choanosomal mesh and zygoses of desmas are similar to those of Gastrophanella implexa from the West Indies. Its tylostrongyles and desmas are similar to those of G. mammilliformis from South Africa. The per- pendicular arrangement of the tylostrongyles, the outwardly directed tyles, the choanosomal desma, and the tylostrongyle type all resemble those of Siphoni- dium capitatum from the Indo-Pacific. Gastrophanella primore and Corallistes isabela (Corallistidae), the only lithistid previously recorded from the central East Pacific, lack common morphological characteristics. There have been few studies of the sponge fauna of the central East Pacific. Early studies near the present collection site do not report any lithistid and are old or incomplete de- scriptions (Carter 1882, Wilson 1904, Dick- inson 1945, Hofknecht 1978). More recently, Green & G6mez (1986), Gomez & Bakus (1992) and Hidalgo (1994) described several species in more detail. A search for records of lithistids from the Mexican Pacific (Gulf of California 31°42'N, 114°48’'W to the Isth- mus of Tehuantepec 14°30’N, 92°10'W, Fig. 1), as well as published studies of areas to the north and south of Mexico, have found only one lithistid sponge, Corallistes isabela (Corallistidae) from the Galapagos Islands at 78 m (Desqueyroux-Fatndez & van Soest 1997). I presently describe a new species of the genus Gastrophanella (Demospongiae: Lithistida: Siphonidiidae) from the Mexican coast, central East Pacific. Materials and Methods The lithistid sponge was dredged by the R/ V El Puma in April 1982, at Guerrero, Mex- ico (Fig. 1). Thirty spicules were measured (in »m) for the mean size and ranges of each type. For scanning electron microscopy (SEM) of spicules, sponge tissue was digest- ed in boiling nitric acid containing glass pearls and was subsequently washed and cen- trifuged (80 g) twice. Two drops of the re- sulting liquid were dried on a glass slide, placed on a stub and sputter coated with gold paladium (300 A thick). Spicules were ob- served with a JEOL JSM-35 scanning elec- tron microscope. The holotype (USNM 51303) is deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Class Demospongiae Sollas, 1888 Order Lithistida Schmidt, 1870 Order used for convenience as it is considered to be polyphyletic (van Soest & Stentoft 1988). Family Siphonidiidae Sollas, 1888 Genus Gastrophanella Schmidt, 1879 Definition.—(Sub)tylostrongyles and (sub)tylostyles arranged perpendicularly to the surface (van Soest & Stentoft 1988). VOLUME 111, NUMBER 4 Mexico East Pacific Fig. 1. Gastrophanella primore, new species Figs. 2A-I, 3 Material examined.—Holotype (USNM 51303), Guerrero, Mexico (16°15'24’N, 98°40'W). Description.—Cup like, 3 cm high by 3.5 cm wide with an apical oval cavity that constitutes the atrium, 10 mm long by 4 mm wide by 14 mm deep, provided with a diaphragm 9 mm below the rim (Fig. 2A, B). Color: grey when alive, nearly white in spirit. Consistency hard; surface apparently smooth, rough to the touch. Oscula on the surface 500 pm in diameter and inside the atrium 90-180 pm. Barely visible ectoso- mal membrane with ostia 2.5 wm in diam- eter. Ectosomal skeleton (Fig. 2C, D) sieve- like with strong zygosed desmas, mesh siz- es 62—127.5 wm in diameter, with monax- ons (sub)tylostrongyles or (sub)tylostyles perpendicular to the surface, loosely inter- spersed among the desmas and with their tyles directed outwardly, reaching, at the base of the sponge, 11.5 wm beyond the surface. Choanosomal desma skeleton (Fig. 2E) with more tightly packed mesh than the VS Gulf of Mexico Guerrero isthmus of Tehuantepec 100 Map showing collection site, Guerrero, Mexico, central East Pacific. ectosomal skeleton; monaxons continue tra- versing the body internally. Spicules.—Desmas rhizoclone (Fig. 2F— H): 147-220.5-274 wm by 12-15-18 pm (minimum-mean-maximum); (sub)tylostyles to (sub)tylostrongyles (Fig. 3) with rugose tyles, occasionally rugose at both ends: 109-326-600 wm by 2.5-5.3-11 wm, tyle di- ameter (Fig. 21) 5.8-8.4-11 pm. Distribution.—So far known only from the type locality, off the coast of Guerrero, Mexico, East Pacific. Etymology.—Named primore as it is the first species of the genus Gastrophanella found in the East Pacific [primore is the feminine of primoris (first) in Latin]. Ecology.—The type locality (Guerrero, Mexico) is characterized by considerable amounts of river runoff due to the outlets of several rivers. Three of the rivers are per- manent, but the remainder are active only during the rainy season. The sponge was collected on a sandy bottom, adhering to fine grained calcareous material at a depth of 45 m. Temperature was 24°C and salinity was 34%o. 776 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Gastrophanella primore, new species: A, holotype habitus, lateral view; B, view from above showing atrium; C, SEM of peripheral skeleton showing ectosomal mesh and oscular morphology (40X); D, SEM of perpendicular section through peripheral skeleton (200); E, SEM of choanosomal desma reticulum (94), note broken perpendicular monaxons; F—H, SEM of desmas rhizoclone (400, 320 and 300%, respectively); I, SEM of enlarged rugose tyle of tylostrongyle (4800X). Scale bars: A, B = 1 cm; C = 200 pm; D = 50 pum; E = 100 pm; F-H = 50 pm; I = 5 pm. VOLUME 111, NUMBER 4 777 Fig. 3. _ Gastrophanella primore, new species: SEM of variations of monaxon: subtylostyles to subtylostron- gyles and tylostrongyles (all same magnification, 2000). Scale bar = 25 ym. Discussion The “order Lithistida” has been recog- nized as an artificial group, designated as incertae sedis by Lévi (1973) and other au- thors. Van Soest & Stentoft (1988) dis- cussed the likely polyphyletic origin of dif- ferent forms of this order, emphasizing the need for a revision of all lithistids. For this reason, the new species, Gastrophanella primore, is assigned only tentatively to the ‘order Lithistida.”’ 778 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Comparisons of G. primore with other species are as follows (see Table 1): Gas- trophanella implexa Schmidt, 1879 sensu van Soest & Stentoft (1988:68), from the West Indies, is the closest species in exter- nal appearance, with its cup shape and tight-meshed choanosomal reticulum, as well as the zygoses of desmas, but differs in the desma shape, spicule bundles and overall measurements. G. mammilliformis Burton (1929:7), from South Africa, agrees in the shape of monaxons and desmas, but the shape of the habitus does not corre- spond to that of the Mexican species, nor does the size of the spicule types and pres- ence of tylostrongyle bundles. Siphonidium capitatum Sollas (1888:317), from the Indo-Pacific, agrees in the perpendicular ty- lostrongyle arrangement and the outward direction of the tyles, its similar choanoso- mal desma and tylostrongyle type, but dif- fers in its fistulose habitus and ramified ec- tosomal desmas, as well as in the diameter of the monaxons. Lithistids occur in deep waters, 100 m to 250 m in average depth. From the most studied regions of the world, the southwest Pacific possesses a rich fauna of about 36 lithistids (taxonomic status unknown): 11 species from Kei Island, Fiji Islands and South Papua at depths of 250 m to 673 m (Sollas 1888), including two shallow-water species at 32 m and 60 m; two species from New Zealand at 108 m and 180 m (Berg- quist 1968); and 23 lithistids from New Caledonia between 175 m and 570 m (Lévi & Lévi 1983, 1988). Similar lithistid diver- sity (32 species) has been reported for the West Indies: 22 species from Cuba, Lesser Antilles and Florida Keys at depths of 100 m to 1450 m (Schmidt 1879, 1880), and 17 valid species, including seven also recorded by Schmidt, from Barbados and other West Indian localities at depths of 100 m to 200 m (van Soest & Stentoft 1988). This con- trasts with the single lithistid Corallistes is- abela Desqueyroux-Fatndez & van Soest 1997, reported in the central East Pacific, even though deep-water sponges have been Depth/bottom type 153/carbonate, biogenic 90-180/not given 45/sand-gravel 182—228/not given 252/blue mud Desmas 280/not given 100/19—27.5 147-274/12-18 150—220/20—25 108/32—60 524/16 480/5 420-570/5 Monaxons 200-395/3-6.5 109—600/2.5—11 Shape Pear, base expanded Tube Cup Fistulose Cup-like Locality Florida Cays Central East Pacific implexa van Soest & Stentoft 1988 Barbados G. ir G. tr G. 1 S. ce G. p South Africa Indo-Pacific Species ammilliformis Burton 1929 pitatum Sollas 1888 more ni. sp. Present work Ss £5 8s eft Table 1.—Comparison of Gastrophanella primore, new species with other species of lithistids (length/diameter measurements in 4m, depth in meters). implexa Schmidt 1879 VOLUME 111, NUMBER 4 collected from the East Pacific: cold tem- perate waters of Southern California, U.S.A. between 0 m and 1865 m (de Lau- benfels 1926, 1932; Bakus & Green 1987; Green & Bakus 1994), the warm temperate Gulf of California in 92 m to 240 m (Dick- inson 1945), and the tropical waters from the Mexican Pacific coast to Central Amer- ica in 95 m to 4433 m (Wilson 1904, Des- queyroux-Fatindez & van Soest 1996). In- deed, further studies of lithistids in this re- gion are needed to determine whether lith- istids are scarce along the central East Pacific. Acknowledgments The author is thankful to Dr. Martin Me- rino for his support and comments, Yolanda Hornelas for the SEM photographs, and Jorge A. Castro Sanchez for the map draw- ing. The comments of anonymous referees improved the original manuscript. Literature Cited Bakus, G. J., & K. D. Green. 1987. The distribution of marine sponges collected from the 1976— 1978 Bureau of Land Management southern California Bight Program.—Bulletin of the Southern California Academy of Science 86(2): 57-88. ; Bergquist, P. 1968. The marine fauna of New Zealand: Porifera, Demospongiae, Part 1 (Tetractinomor- pha and Lithistida)—New Zealand Department of Scientific and Industrial Research Bulletin 188:1—-105. Burton, M. 1929. Descriptions of South African spong- es collected in the South African marine survey. Part II. “Lithistidae,” with a critical survey of the desma-forming sponges.—Union of South Africa Fisheries and Marine Biological Survey, Report 7:1—12. Carter, H. J. 1882. Some sponges from the West Indies and Acapulco in the Liverpool Free Museum described, with general and classificatory re- marks.—Annals and Magazine of Natural His- tory (ser. 5) 9:266—301, 346-368. Desqueyroux-Fatindez, R., & R. W. M. van Soest. 1996. A review of Iophonidae, Myxillidae and Tedaniidae occurring in the South East Pacific (Porifera: Poecilosclerida)—Revue Suisse de Zoologie 103(1):3-79. , & R. W. M. van Soest. 1997. Shallow waters 779 Demosponges of the Galapagos Islands.—Re- vue Suisse de Zoologie 104(2):379—467. Dickinson, M. G. 1945. Sponges of the Gulf of Cali- fornia.—Allan Hancock Pacific Expeditions 11(1):1—251. Gomez, P, & G. J. Bakus. 1992. Aplysina gerardo- greeni and Aplysina aztecus (Porifera: Demo- spongiae), new species from the Mexican Pa- cific.—Anales del Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Aut6noma de México 19(2):175—180. Green, G., & P. Gomez. 1986. Estudio taxonémico de las esponjas de la Bahia de Mazatlan, Sinaloa, Mexico.—Anales del Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Au- tonoma de México 13(3):273—300. Green, K. D., & G. J. Bakus. 1994. Taxonomic atlas of the benthic fauna of the Santa Maria Basin and Western Santa Barbara Channel.—Santa Barbara Museum of Natural History California, 82 pp. Hidalgo, A. 1994. Taxonomia del Phylum Porifera (Clase Demospongiae) del Mar de Cortes.— Tesis Profesional, Universidad Nacional Aut6n- oma de Mexico, 110 pp. Hofknecht, G. 1978. Descriptions and key to the in- tertidal sponges of the Puerto Penasco area in the Gulf of California.—Journal of the Arizona- Nevada Academy of Science 13:51—56. Laubenfels, M. W. de 1926. New sponges from Cali- fornia—Annals and Magazine of Natural His- tory (9)17:567-573. . 1932. The marine and fresh water sponges of California.—Proceedings of the United States National Museum 81, art. 4:1—140. Lévi, C. 1973. Systématique de la classe des Demo- spongiaria (Démosponges). Pp. 576-631 in P. Grassé, ed. Traité de Zoologie: Anatomie, Sys- tématique, Biologie, Masson et C* Editeurs, Paris 3(1):715. . & P. Lévi. 1983. Eponges Téractinellides et Lithistides bathyales de Novelle-Calédonie.— Bulletin de Muséum National d’Histoire Natu- relle 4(5)(A, 1):101—168. , & . 1988. Nouveaux spongiaires lith- istides bathyaux 4 affinités crétacées de la Nou- velle-Calédonie—Bulletin de Muséum National d’ Histoire Naturelle 4(10) (A, 2): 241-263. Schmidt, O. 1970. Grundztige einer spongien-fauna des Atlantischen Gebietes. Leipzig, Engelmann i-iv, 1-88 pp. . 1879. Die spongien des Meerbusen von Mex- ico I. Abt. Lithistiden I. Heft. G. Fischer Jena, 32 pp. 4 pl. . 1880. Die spongien des Meerbusen von Mex- ico (und des Caraibischen Meeres) II. Abt. Hex- actinellida, III. Abt. Tetractinellida, Monacti- 780 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nelliden und Anhang II. Heft. G. Fischer Jena, the Voyage of H. M. S. Challenger, Zoology 35—90, 5-10 pl. 25(63):i—clxvi, 1-458 pp., 44 pl. Soest, R. W. M. van, & N. Stentoft. 1988. Barbados deep- Wilson, H. V. 1904. The sponges. Reports on an ex- water sponges.—Studies on the Fauna of Curacao ploration off the West coasts of Mexico, Central and other Caribbean Islands 70(122):175. and South America, and off the Galapagos Is- Sollas, W. J. 1888. Report on the Tetractinellida col- lands.—Memoirs of the Museum of Compara- lected by H. M. S. Challenger during the years tive Zoology 30(1):1—164. 1873—76.—Report on the Scientific Results of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):781—794. 1998. Asexual reproduction in Linuche unguiculata (Swartz, 1788) (Scyphozoa: Coronatae) by planuloid formation through strobilation and segmentation Fabio Lang da Silveira and André Carrara Morandini Departamento de Zoologia, Instituto de Biociéncias, Universidade de Sao Paulo, Caixa Postal 11461, 05422-970 Sao Paulo, SP, Brazil Abstract.—Scyphistomae of Linuche unguiculata from Sao Paulo State, Bra- zil, were reared for one year to study the life cycle of this warm-water species (in subtropical western South Atlantic waters). We found L. unguiculata re- produces more by the development of planuloids than by medusae. The scy- phistoma is rejuvenated by an operculated regression-regeneration cycle. We hypothesize that reproduction of the species by production, either via segmen- tation or strobilation, and liberation of planuloids explains the absence of re- ports of ephyrae and medusae in the area. Jarms (1997) reviewed the history of knowledge about scyphistomae of Corona- tae. The only records of coronate Scypho- zoa from the Brazilian coast are of the me- dusa stage of Nausithoe punctata KOolliker, 1853, from the north of Bahia State (Goy 1979) and material recorded during study of the life cycle of Nausithoe aurea Silveira & Morandini, 1997, from the north of Sao Paulo State. Silveira & Morandini (1996) redescribed the scyphistoma of a coronate, Stephanoscyphistoma corniformis (Komai, 1936), from the south-east Brazilian coast. The present study on the life cycle of L. unguiculata was undertaken over a 14 month interval. Observations were based on cultures of the scyphistomae from south- east Brazil. We found colonial coronate scyphisto- mae (presumed to be L. unguiculata) fre- quently in Sao Sebastiao Channel, and we asked: why are there no records of medusae or of any dermatitis caused by their planu- lae? We presumed three possible explana- tions to account for this: the ephyrae belong to suprabenthic communities and are trans- ported to distant places, so that medusae of this species are not found near the coast; L. unguiculata typically passes intervals of many years between periods of production of large numbers of medusae (Russell & Tomchik 1993, Black et al. 1994); therefore it would not be strange that between these periods the probability of finding medusae were very small; the species was not L. un- guiculata, although the characters of the periderm tubes of our material were similar to that reported in the species. Our field observations were that no ephy- rae/medusae occurred in plankton or supra- benthic samples during 1 yr and our labo- ratory observations were that few polyps strobilated producing a small number of ephyrae, but many planuloids were pro- duced together with segmentation of other scyphistomae. Thus, few ephyrae were pro- duced, which supports the second explana- tion. Material and Methods Colonial scyphistomae (Fig. 1) were sampled from calcareous debris, mainly fragments of the stony coral Mussismilia hispida (Verrill, 1902) (Scleractinia, Mus- sidae) at 2-6 m depth by SCUBA diving, in Sao Sebastido Channel (23°50’S, 782 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Colony of Linuche unguiculata, as seen soon after sampling. The living tissues are much contracted within the base of the tubes. Note that the scyphorhiza (arrows) is delicate and encrusting. Scale: 0.6 mm. VOLUME 111, NUMBER 4 783 Iie, Ze. Scale: 1.25 mm. Colony of Linuche unguiculata. The oral 45°25'W), of the rocky shore at Urubu Point and on Praia Grande reef, Ilhabela County, SP. Sampling was done on one or two days of every month, except August, from June 1996 until May 1997. We searched for the colonies on lumps of the calcareous substratum collected, with the aid of a stereomicroscope. Each monthly sample included a minimum of 15 colonies, discs of the scyphistomae are partly or fully extended. each with 2 to 8 living polyps (Fig. 2). The colonies were transferred into and main- tained in small Petri dishes containing about 70 ml of filtered sea-water. At least twice a month, the colonies were cleaned by gentle rubbing with a delicate brush or by removing the many filamentous algae with fine forceps. In each search for medusae or ephyrae, 784 three plankton and suprabenthic faunal samples were taken on three consecutive days following the sampling of calcareous debris. Twenty-minute tows were made in the vicinity of the sampling sites of the de- bris with a plankton net and a dredge (the last equipment according to Wakabara et al. 1993) at a depth of 5 to 10 m, both with 0.5 mm stretch mesh, using a small out- board motor boat. Two divers followed the equipment to maintain the plankton net at 3 m below the surface and to prevent the dredge from being clogged with bottom sediments. The scyphistomae reared in the Centro de Biologia Marinha da Universidade de Sao Paulo (CEBIMar USP), at Sao Sebastiao, SP, were treated as follows. From 6 June to 25 July 1996 all colonies were kept at room temperature (18°C—28°C) and from 1 April to 7 June 1997 half were kept at room tem- perature (21°C—30°C, although during 24 h the temperature range never exceeded 4°C) and one-half were treated with temperature changes, 18—27—18°C (based on the month- ly averages of surface water temperature for winter and summer in the area), at 7-day intervals and /,, h light/dark regime inside an incubator (FANEM® 347-CDG). The scyphistomae reared in the Zoology De- partment (IB, USP), at Sao Paulo, SP, were treated as follows. From 26 July 1996 to 11 June 1997 the June—February samples were kept at controlled temperature (August—mid February a monthly increase of 1°C starting at 21°C; mid February—June, a monthly de- crease of 1°C starting at 26°C—following the natural surface water temperature monthly averages) and %, h light/dark re- gime inside an incubator. The sea water was changed and the animals were fed every other day with a deep-frozen stock of ho- mogenate of the clam Perna perna (Lin- naeus, 1767) (Bivalvia, Mytilidae). Some of the specimens collected in April 1997 were used to measure the periderm tube (total length, diameter at aperture and base), and the internal cusp was examined in 42 pol- yPps. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Free-swimming ephyrae, from strobilat- ing scyphistomae, were transferred into sin- gle airtight pots (universal samplers for medical analysis ZESTER® 140 ml). They were fed from a deep-frozen stock of ho- mogenized clam gonads. The ephyrae and medusa were fed every day. They were in- dividually immersed in the homogenized food, observed under a stereomicroscope until their stomachs were full, and removed to a new pot with clean filtered sea water. Free-swimming planuloids (samples of March, April and May 1997), produced by strobilating or from segmenting scyphisto- mae, were transferred into single dishes, at 24 h intervals, for further observations. We studied the cnidome of live speci- mens and of specimens preserved in sea- water-formaldehyde solution. In preserved tissues, only undischarged nematocysts were measured. Voucher specimens are: National Muse- um of Natural History, Smithsonian Insti- tution (USNM 99376, 99377); Museu de Zoologia da Universidade de Sao Paulo, Brazil (MZUSP 12.355, 12.356); Museu Nacional Universidade Federal do Rio de Janeiro, Brazil (MNRJ 3126, 3127, 3128); The Royal Ontario Museum, Canada, In- vertebrate Zoology (ROMIZ B3010, B3011); The Zodlogisch Museum, Univer- sity of Amsterdam, The Netherlands, (ZMA Coel. 8507, 8508); Zodlogisches Museum, Universitat Hamburg, Germany, (ZMH C11639, C11640, C11641); and The Natu- ral History Museum, United Kingdom, (NHM 1998.145, 1998.146). Results We found no ephyrae or medusae of Lin- uche unguiculata in plankton and supra- benthic samples. During June 1996—June 1997, over 240 scyphistomae were reared in the Zoology Department, at Sao Paulo City, but none of them strobilated. Never- theless, on many occasions we observed that some scyphistomae produced a peri- derm operculum closing the tube. The oral VOLUME 111, NUMBER 4 disc and the gastric septa of those scyphis- tomae regressed, but the column remained and they were able to contract and extend within the tube. We followed some closed tubes over a varied period of 1—3 d, at the end of which the operculum opened and the polyp regenerated septa and the oral disc. During the period late March—June 1997, among 559 scyphistomae (some 130 colo- nies) reared at room temperature in CEBI- Mar, only 6 strobilated (two in the same colony) and 3 segmented irregularly. At the start of strobilation, the polyp re- sorbed the oral disc, the strobilating region occurred over most of the column, disc number varied from 8 to 17, and no oper- culum, of either periderm or tissue was present. Initially, the discs were set well apart and the remaining column tissues be- tween them were bulged. From a few hours to 3 d, the discs changed into irregular tis- sue fragments and most of the remaining and interconnecting column tissues con- stricted. Only the distal disc enlarged and metamorphosed into an ephyra (Fig. 3), while the other discs directly transformed into planuloids. Among the six strobilae, five released one ephyra each and in anoth- er the ephyra transformed into a planuloid within the tube. Release of the ephyra (Fig. 4) varied from 1 to 3 d after onset of stro- bilation (Table 1). The earliest stage at which the ephyra acquired noticeably me- dusa-like characters was at about 5 mm in diameter (specimen 8, Table 1). The gonad primordia were four pairs of radiating tissue cords on each side of the perradii. At about 47 d, each gonad pair had fused giving rise to four cleft crescents (Fig. 5). This medusa had an anomalous number of lappets due to an injury at early ephyra stage (fourteen lappets, two perradial pairs being fused, Fig. 5). Most of the zooxanthellae were ir- regularly distributed within the gastroder- mis, in side view mostly below the coronal groove (Fig. 5) and in the corners of the lips of the manubrium. The lappets were round, and slightly overlapping in an or- derly fashion, which made the medusa ro- 785 tate clockwise upon its oral-aboral axis while swimming. The short, translucent ten- tacles were situated within the clefts be- tween the lappets. The eight conspicuous rhopalia were directed outward while the medusa was at rest or downward during contraction of the bell. The ephyrae of the remaining material were preserved before reaching the medusa stage (Table 1). At the start of segmentation, the polyp resorbed the oral disc and produced a peri- derm operculum, it acquired a transverse segmenting region over most of the column, and the segments tended to become spher- ical in shape. In one colony we observed that in one operculate scyphistoma many planuloids appeared by irregular segmen- tation. These fused together to restore the column of the polyp, but without differen- tiation of septa and oral disc (see sequence in Figs. 6, 7). Thus, the process lasted until the end of April. In some colonies, we not- ed that after a few weeks, near the oper- culum margin, a fissure developed through which a few planuloids could emerge (Fig. 8). The crack was apparently caused by the egress of the planuloids. The measurements, length of the tubes and diameter at the base and at the aperture, are: 1.8—12.7 mm, 0.12—0.49 and 0.25—0.68 mm. Most tubes had one cusp; none, two or three were less frequent. The cusps were cup-shaped, with longitudinal ridges, main- ly at the broad round margin (Fig. 9). The axis of the cusp base outline was long, nar- row, and parallel with the long axis of the tube, whereas the proximal end was bulging and circular (Fig. 9). Liberated planuloids were elliptical, swam actively by cilia, and had transparent epidermis and a gastrodermis with many zooxanthellae. Two planuloids (from spec- imen 8, Table 1) settled on the bottom of a small Petri dish on 27 May and O1 June respectively and were reared for 58 and 54 d. Each zooxanthellate polyp had five ten- tacles within 3 d. Each produced a delicate cylindrical tube, without branching and without cusps. 786 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Strobila of Linuche unguiculata (April 1997). A, early strobilation phase, at 24 h, in which the connections between the discs narrow and the distal disc is bigger; B, late strobilation phase, at 72 h, in which the distal disc has differentiated into an ephyra and the remaining ones into planuloids. Scale: 0.6 mm. VOLUME 111, NUMBER 4 787 +34 5 nb MH, th es 3 a 4 e gh aua€ ~~ Pig “Se 2 so ee * Fig. 4. Ephyra (oral view) of Linuche unguiculata, two days after release from the strobila. Note the many zooxanthellae. (From photomicrograph). Abbreviations: cg = coronal groove, gf = gastric filament, r = rho- palium, z = zooxanthellae. Scale: 0.6 mm. Scyphistomae, planuloids and ephyrae (Table 2) all had holotrichous isorhizas and heterotrichous microbasic euryteles nema- tocysts. Discussion Our original assumption was that the co- lonial coronates in Sao Sebastiao Channel were Linuche unguiculata, considering the morphology and number of the internal cusps within the tube. The cup-shaped cusp is diagnostic of polyps of L. unguiculata (Leloup 1937, Ortiz-Corp’s et al. 1987, Jarms 1991). Moreover, the size and pro- portions of the periderm tubes were within the ranges of L. unguiculata studied by Le- loup (1937), Allwein (1968) and Ortiz- Corp’s et al. (1987). The original descrip- tion of the species was based on the medusa stage (Swartz 1788) and it is wise to study both stages in the life-cycle to make a pre- cise identification (Jarms 1990, 1991). Wer- ner (1979) was first to link the polyp stage of Stephanoscyphus komaii with the me- dusa L. unguiculata. The lack of either ephyrae or medusae in plankton and suprabenthic faunal samples had been difficult to explain. Their absence was evidently due to the asexual reproduc- tive modes employed locally—direct trans- formation of strobilating discs and segmen- tation originating planuloids, newly report- ed reproductive strategies for L. unguicu- lata and novelties for coronates. Arai (1997:166) defined strobilation as the process by which scyphistoma produces 788 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Observations on 8 colonial scyphistomae of Linuche unguiculata. Abbreviations: St-e—length of period of strobilation (ds) originating ephyrae, St-p—tength of period of strobilation (ds) originating planuloids, Se—length of period of segmentation (ds), E—ephyra development (ds), E/M—observations (ds) of ephyrae and medusa. Scyphistoma 1 2 3 4 S) 6 7 8 (sample month) (Jun 96) (Mar 97) (Mar 97) (Mar 97) (Mar 97) (Mar 97) (Mar 97) (Apr 97) St-e 3 2 and 1 — — — — 1 2 St-p 7 8 and 1 — — 7 — 5 a Se — — 8 7 — 10 — — E = = ats ae aus = = 30 E/M 36 24 and 38 — — — — Di 54 2 The ephyra was preserved to study the cnidome. ephyrae: ‘‘This requires disc formation (‘segmentation’) leading to fission, and also metamorphosis in which structures of the polyp are lost and replaced in each disc with those of the developing ephyrae.”’ Therefore, the word segmentation in this work applies to the transverse fission of the scyphistoma, within an operculate tube, comparable with the strobilation in which all discs metamorphose into ephyrae. Wer- GRCM Fig. 5. Young medusa of Linuche unguiculata. The animal is shown in side view, to show the dark pattern due to the zooxanthellae in the gastrodermis and the arrangement of the paired gonads in cleft crescents. Note that the nearest lappet is enlarged as result of injury and later fusion of two original lappets. (From life and close-up VHS). Scale: 1 mm. VOLUME 111, NUMBER 4 789 GRC M GRCM CINE IM GRUY\ Fig. 6. Colony of Linuche unguiculata undergoing segmentation. A, planuloid formation (3.1V.97); B, plan- uloids almost distinct and the distal ones fused together (4.1V.97); C, regeneration of the column of the polyp starts by fusion of planuloids (dark areas) and the stretching of basal tissues (9.1V.97); D, contraction of the basal tissues, upon stimulation with a forceps, showing that the distal planuloids are entirely fused together (9.I1V.97). Scale: 1.25 mm. ner (1973) reviewed the known variation of the operculum in the scyphistomae of Co- ronatae. The only paper with a thorough ac- count of the strobilation of L. unguiculata is that of Ortiz-Corp’s et al. (1987). They observed the process in Puerto Rico, at temperatures of 25—28°C (December—May) and 28—30°C (June—November), for 7 yr. They noted strobilation once a year, some- times within an operculate tube, with the production of free-swimming ephyrae. Wer- ner (1979) also accounted for the annual strobilation of L. unguiculata under labo- ratory conditions. Werner (1979:94) hy- pothesized that for the colonial coronates “... their strobilation activities are geneti- cally fixed to one distinct short season.’ He argued that the surplus of energy stored by the polyps is available for colony growth, in place of consecutive strobilation within one year the way solitary coronates are likely to do. Jarms (1997) reported shortened life cy- cles of five species of Coronatae, the trans- formation of free ephyrae into planuloids and these into new polyps, due to unfavor- able conditions of salinity, temperature or food supply. Planuloid formation by trans- formation of free ephyrae within the tube is known in the solitary Nausithoe planulo- phora (Werner 1971, Werner & Hentschel 1983, Jarms 1997) and N. aurea (Silveira & Morandini 1997), and by parthenogene- sis in Thecoscyphus zibrowii (Werner 1984). A unique feature we have observed is the differing destiny of discs from the same strobila—one ephyra and many plan- uloids—with differing regulative mecha- 790 GAGM Fig. 7. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ess ii Sy C' Colony of Linuche unguiculata undergoing segmentation. A, the column of the polyp is almost entirely regenerated (13.1V.97); B, contraction of the tissues, upon stimulation with a forceps, to show that there is integration between the regenerated pieces, but that the column as a whole does not contract (13.1V.97); C, column totally regenerated, but the dark areas indicate the remains of fusing planuloids (14.IV.97). Scale: 1.25 nm. nisms for each developing structure. Fol- lowing the reasoning of Jarms (1997:275) on planuloid formation by N. planulophora, we believe the planuloids of L. unguiculata must be an alternative dispersive stage, since we have observed settlement and de- velopment of two new colonies. We hy- pothesize that segmentation is an advanced trait derived from the peculiar strobilation, with the addition of the operculum, perhaps the most derived response to the regulation of asexual reproduction in the species. L. unguiculata is a warm-water species (May- er 1910, Kramp 1961, Ortiz-Corp’s et al. 1987), and we have observed a new repro- ductive mechanism of the species to adapt to a subtropical area in which there is a marked influence of both warm and cold water masses (see Campaner 1985). The ab- sence of ephyra and medusa from plankton and suprabenthic samples is thereby ex- plained. The ephyrae (Fig. 4) and the young me- dusa (Fig. 5) closely resemble the descrip- tions of the adult medusa of L. unguiculata (Mayer 1910, Ortiz-Corp’s et al. 1987). The medusa illustrated in the present paper re- sembles the developmental stage that May- er (1910:559) described “‘that the gonads appear when the medusa is about 5 mm wide.”’ The arrangement of the gonads and the shape and coloration of the umbrella are VOLUME 111, NUMBER 4 791 Fig. 8. Photomicrograph of the scyphistoma periderm tube of Linuche unguiculata, distal end and with operculum (white arrow). Note that one planuloid (black arrow) has emerged and rests upon the operculum. Scale: 0.3 mm. Table 2.—Measurements of cnidae of cnidome of Linuche unguiculata. The range was taken from 20 nematocysts of each type at each stage, except the un- derlined measurements taken from only 9 nematocysts. Holotrichous isorhiza Heterotrichous microbasic (wm) eurytele (4m) Scyphistoma 10-6 X 6-4 17-14 X 14-11 13-07 X 10-05 Ephyra 7-5 x4 15-14 Xx 14-12 13-09 X 12-08 Planuloid 7-6 X 5-4 17-14 X 14-12 13-09 X 12-07 diagnostic for the species. The difficulty of obtaining the strobilae and the ephyrae-me- dusae reflects the prevailing asexual repro- duction. The cnidome of scyphistoma, ephyra and planuloid are the same as re- corded by Calder (1974) and Ortiz-Corp’s et al. (1987) in medusae of L. unguiculata. The differences between our data may re- sult from comparing different stages for the species. We followed Calder (1974) in sep- arating the heterotrichous microbasic eury- tele nematocysts into two size-classes, though they are almost one continuous size class. There are some indirect indications, e1- 792 GRCM Fig. 9. Segment of periderm tube of Linuche un- guiculata showing an internal cusp (since some had 2— 3). Arrow points to the base of the outline of the cusp. Note that at the broad round margin (M) there are ridg- es. Scale: 0.3 mm. ther in the swarming of the medusa (Larson 1982, Ortiz-Corp’s et al. 1987, Halstead 1988), as well as in the outbreaks of their planulae (Russell & Tomchik 1993, Black et al. 1994), that in nature L. unguiculata strobilates seasonally (winter—spring) and the adult medusae occur in spring and early summer in the Caribbean, Atlantic coast of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Florida, and the Bahamas. Although some authors report concentrations of medusae of L. unguiculata (Table 3), only two medical incidents (dermatitis) linked with the me- dusae of this species have been reported in the literature (Penner 1962). Black et al. (1994) associated seabather’s eruption— SBE—with planulae of L. unguiculata in Florida. Russell & Tomchik (1993) and Black et al. (1994) summarized outbreaks of SBE in Florida. They argued that there may be an interval of up to 25 yr, between major outbreaks. Nevertheless, Williamson et al. (1996:308—310) stated that the der- matitis SBE, besides having other names, is caused by different organisms in distinct re- gions. We hypothesize that L. unguiculata is a species with a long life-cycle mainly resulting from the long duration of the scy- phistoma stage. Under variable conditions, this warm-water species may present an ef- ficient colonizing ability by the scyphisto- ma that undergoes asexual reproduction and propagation. The oral disc regression-regeneration process we observed in operculate scyphis- tomae on many occasions has never been reported in any coronate. This mechanism may be considered analogous to observa- tions on some leptolid polyps, e.g., Thyros- cyphus marginatus (Harris 1990, and see revision in Crowell 1991). Tardent (1963) hypothesized that a regression-regeneration cycle in the hydranths of colonial leptolids would be a natural histophysiological reju- venation mechanism for the colony, by ex- tension a possible pattern for some species to secure perennation (sensu Cornelius 1992) in specific habitats. Nevertheless, it Table 3.—Occurrence of swarms of medusae of Linuche unguiculata in Florida, the Caribbean and the Ba- hamas, from different sources. Occurrence Date Miami (Florida) April 1950 Sarasota (Florida) July 1954 Boca Grande (Florida) July 1955 Carrie Bow Cay (Belize) The Bahamas April/May 1989 Mar/Apr 1987; April 1988; Mar/Apr 1990 Author Penner 1962 Penner 1962 Penner 1962 Larson 1992 Montgomery & Kremer 1995 VOLUME 111, NUMBER 4 remains an open question whether this pro- cess in laboratory observations really oc- curs under natural conditions (see revision in Hughes 1987). Acknowledgments We thank the Director and staff of CE- BIMar, at Sao Sebastiao, SP, for their sup- port to accomplish this work. We are deeply indebted to Dr. A. E. Migotto (CEBIMar USP) for helping us to solve some technical difficulties. We are indebted to Dr. G. Jarms (Zoologisches Institut und Zoologisches Museum der Universitat Hamburg), to Dr. Dale Calder (The Royal Ontario Museum) and to an anonymous reviewer for the re- visions of the manuscript. We are also in- debted to Dr. P. E S. Cornelius (The Natural History Museum, London) for revision of the manuscript and of the English version. We thank Drs. Y. Wakabara and A. Tararan (IO USP) for lending their suprabenthos dredge at the start of the research. We are deeply indebted to G. R. C. Monteiro (FITO) for the line art drawings. This work was supported by FAPESP, grants 96/1416-9 and 96/1415-2 and USP, grant 96/13802612. Literature Cited Allwein, J. H. 1968. Stephanoscyphus komaii Leloup (Scyphozoa, Coronatae) from the Caribbean.— Videnskabelige Meddelelser fra dansk naturhis- torisk Forening 1 Kobenhavn 131:193—197. Arai, M. N. 1997. A functional biology of Scyphozoa. Chapman & Hall, London, xvi, 316 pp. Black, N. A., A. M. Szmant, & R. S. Tomchik. 1994. Planulae of the scyphomedusa Linuche ungui- culata as a possible cause of seabather’s erup- tion.—Bulletin of Marine Science 54(3):955— 960. Calder, D. R. 1974. Nematocysts of the coronate scy- phomedusa, Linuche unguiculata, with a brief reexamination of Scyphozoa nematocyst clas- sification.—Chesapeake Science 15:170—173. Campaner, A. FE 1985. Occurrence and distribution of copepods (Crustacea) in the epipelagial off southern Brazil.—Boletim do Instituto Ocean- ografico, Universidade de Sao Paulo 33(1):5— Dale Cornelius, P FE S. 1992. Medusa loss in leptolid Cni- 793 daria, hydroid rafting, and abbreviated life-cy- cles among remote-island forms: an interim re- view.—Scientia Marina 56(%):245—261. Crowell, S. 1991. Regression and replacement of hy- drants in thecate hydroids, and the structure of hydrothecae.—Hydrobiologia 216—217:69-73. Goy, J. 1979. Campagne de la Calypso au large des cdtes atlantiques de 1’ Amérique du Sud (1961— 1962)—35.—Meéduses. Résumé scientifique de la campagne de la Calypso au large des cétes atlantiques de 1’ Amérique du Sud (1961-1962) 11:263-296. Halstead, B. W. 1988. Chap. 4—Invertebrates, Phylum Coelenterata (Cnidaria). Pp. 99-186 in Poison- ous and venomous marine animals of the world. 2nd edition. The Darwin Press Inc., Princenton, 288 pp. Harris, V. A. 1990. 11. Hydroids (Cnidaria: Hydrozoa). Pp. 223-269 in Sessile animals of the sea shore. Chapman & Hall, London, 379 pp. Hughes, R. G. 1987. 16. The loss of hydranths of La- omedea flexuosa Alder and other hydroids, with reference to hydroid senescence Pp. 171-184 in J. Bouillon, E Boero, FE Cicogna, & P. ES. Cor- nelius, eds., Modern trends in the systematics, ecology and evolution of hydroids and hydro- medusae. Oxford University Press, Oxford, 328 Pp: Jarms, G. 1990. Neubeschreibung dreier Arten der Gattung Nausithoe (Coronata, Scyphozoa) sow- ie Wiederbeschreibung der Art Nausithoe mar- ginata Kolliker, 1853—Mitteilungen aus dem hamburgischen zoologischen Museum und In- stitut 87:7—39. . 1991. Taxonomic characters from the polyp tubes of coronate medusae (Scyphozoa, Coron- atae).—Hydrobiologia 216—217:463—470. . 1997. The polyps of Coronatae (Scyphozoa), a review and some new results. Pp. 271—278 in J. C. den Hartog, ed., Proceedings of the 6th International Conference on Coelenterate Biol- ogy 1995. Nationaal Natuurhistorisch Museum, Leiden, xviii, 542 pp. Kramp, P. L. 1961. Synopsis of the medusae of the world.—Journal of the Marine Biological As- sociation of The United Kingdom 40:1—469. Larson, R. J. 1982. Medusae (Cnidaria) from Carrie Bow Cay, Belize.—Smithsonian contributions to the Marine Sciences 12:475—488. 1992. Riding Langmuir circulations and swimming in circles: a novel form of clustering behavior by the scyphomedusa Linuche ungui- culata.—Marine Biology 112:229—235. Leloup, E. 1937. Hydropolypes et scyphopolypes re- cueillis par C. Dawydoff sur les cétes de l’Indochine frangaise. II. Scyphopolypes.—Me- moires de Musée Royal d’ Histoire Naturelle de Belgique, Deuxiéme Série 12 12:59-—70. 794 Mayer, A. G. 1910. Medusae of the World. Volume III. The scyphomedusae. Carnegie Institution, Washington D.C., pp. 499-735. Montgomery, M. K., & P. M. Kremer. 1995. Trans- mission of symbiotic dinoflagellates through the sexual cycle of the host scyphozoan Linuche unguiculata.—Marine Biology 124:147—155. Ortiz—Corp’s, E., C. E. Cutress, & B. M. Cutress. 1987. Life history of the coronate scyphozoan Linu- che unguiculata (Swartz, 1788).—Caribbean Journal of Science 23:432—443. Penner, L. R. 1962. Stinging due to Linuche unguicu- lata (Schwartz) in Florida waters.—Quarterly Journal of the Florida Academy of Sciences 25(2):121—-126. Russell, M. T., & R. S. Tomchik 1993. Seabather’s eruption, or “sea lice’: new findings and clin- ical implications.—Journal of Emergency Nurs- ing 19(3):197—201. Silveira, EF L. da, & A. C. Morandini, 1996. Stephan- oscyphistoma corniformis (Komai, 1936) (Cni- daria, Scyphozoa, Coronatae) from the north coast of S40 Paulo, Brazil—SEMINA, Ciéncias Bioldgicas/Satide 17(2):137—145. . 1997. Nausithoe aurea n. sp. (Scyphozoa, Co- ronatae, Nausithoidae), a species with two path- ways of reproduction after strobilation: sexual and asexual.—Contributions to Zoology 66(4): 235-246. Swartz, O. 1788. Medusa unguiculata och Actinia pus- illa.—Kungliga Svenska Vetenskaps Akade- miens Handlingar 9:198—202. Tardent, P. 1963. Regeneration in the Hydrozoa.—Bi- ological Review 38:293-333. Wakabara, Y., A. Tararam, & M. N. Flynn. 1993. Im- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON portance of the macrofauna for the feeding of young fish species from infralittoral of Arroz- al—Cananéia lagoon estuarine region (25°02'S— 47°56'’W)—Brazil.—Boletim do _ Instituto Oceanografico, Universidade de Sao Paulo 41(%4):39-S2. Werner, B. 1971. Stephanoscyphus planulophorus n.spec., eine neuer Scyphopolyp mit einem neuen Entwicklungsmodus.—Helgolaender wissenschaftliche Meeresuntersuchungen 22: 120-140. . 1973. New investigations on systematics and evolution of the class Scyphozoa and the phy- lum Cnidaria.—Publications of the Seto Marine Biological Laboratory 20:35-61. . 1979. Coloniality in the Scyphozoa: Cnidaria. Pp. 81-103 in G. Larwood, & B. R. Rosen, eds., Biology and systematics of colonial organ- isms. Academic Press, London, 589 pp. . 1984. Stamm Cnidaria, Nesseltiere. Pp. 11— 305 in H.-E. Gruner, ed., Lehrbuch der Spe- ziellen Zoologie (coordinated by A. Kaestner), Band I, Wirbellosee Tiere, 2 Teil, Cnidaria, Ctenophora, Mesozoa, Plathelminthes, Nemer- tini, Entoprocta, Nemathelminthes, Priapulida. G. Fisher, Stuttgart, 621 pp. , & J. Hentschel, 1983. Apogamous life cycle of Stephanoscyphus planulophorus.—Marine Biology 74:301—304. Williamson, J. A., P. J. Fenner, J. W. Burnett, & J. E Rifkin, eds., 1996. Venomous and poisonous marine animals: a medical and biological hand- book. Brisbane, Queensland: Surf life Saving Queensland Inc. Kensington, NSW: University of New South Wales Press Ltd., 580 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):795-798. 1998. Acteonina permiana, a new species from the Permian Coyote Butte Formation, central Oregon (Mollusca: Gastropoda: Actaeonidae) Rex Alan Hanger and Ellen E. Strong (RAH) Department of Geology, George Washington University, Washington, D.C. 20052, U.S.A.; (EES) Department of Biological Sciences, George Washington University, Washington, D.C. 20052, U.S.A. Abstract.—Acteonina permiana, is described from the Early Permian Coy- ote Butte Formation of Crook County, central Oregon. Occurrence of Ac- teonina indicates affinity with the Carboniferous to Permian Boreal faunal province, and implies that the Grindstone terrane need not have an “‘exotic”’ origin thousands of kilometers from the North American continent during the Permian. Acteonina is among the rarest of all Up- per Paleozoic gastropod genera. Knight (1932, 1936, 1941) counted only two known specimens, both from the Visean of Belgium. In their survey of Paleozoic opis- thobranch gastropods, Kollmann & Yoch- elson (1976) point out that the genus, Ac- teonina, “‘... has not yet been found in North America in spite of careful search among large collections of Pennsylvanian and Permian gastropods.”’ Acid etching of limestone samples from the Early Permian (Wolfcampian-Leonardina) Coyote Butte Formation of central Oregon has produced many new taxa (Hengstenberg et al. 1997), including four specimens of the new spe- cies, Acteonina permiana. The presence of Acteonina in Oregon has important implications for western North American terrane paleogeography. The Coyote Butte Formation is part of the Upper Paleozoic—Mesozoic Grindstone ter- rane of central Oregon (Wardlaw et al. 1982), and occurs as chaotically inter- mixed limestone blocks within cherts and volcaniclastics. The limestone is interpret- ed as slide and slump blocks that became detached from a carbonate shelf and incor- porated into deeper-water basinal clastics in a forearc basin (Blome & Nestell 1991). The exact dimensions of this basin, and specifically the longitudinal separation of this island arc from the North American mainland remain controversial, and dis- tances of greater than 5000 km with south- ern hemisphere origins have been hypoth- esized (Jones 1990, Miller et al. 1992). The Coyote Butte Formation fauna has been only partially described: brachiopods (Cooper 1957); fusulinids (Skinner & Wilde 1966), colonial rugose corals (Ste- vens & Rycerski 1983) and conodonts (Wardlaw et al. 1982). Acid etching reveals the presence of many undescribed gastro- pods, bivalves, chitons, rostroconchs, nau- tiloids, bryozoans, sponges, trilobites, os- tracods and annelids. A detailed paleobi- ogeography, incorporating the paleoecolog- ical framework, is desirable, but much basic taxonomic work needs to be completed on these important taxa. Of the described fauna, the fusulinids and corals have thus far been most useful for paleobiogeographic comparison. The colo- nial rugose corals are part of a “Thysano- phyllum Belt” fauna that existed along the northern and western margins of Pangea, but Stevens and Rycerski (1983) suggested 796 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. Holotype USNM 498736. Bar is 1 mm; 2, Holotype USNM 498736, close-up of protoconch, showing heter- ostrophic larval whorl. Bar is 100 wm; 3, Paratype USNM 498737. Bar is 1 mm. that the Coyote Butte fauna might have ex- isted several hundred kilometers west of the margin based upon high Otsuka coefficient similarity with known exotic terranes. Fu- sulinids suggest a position at equatorial lat- itudes and many hundreds of kilometers west of the Permian North American mar- gin (Ross and Ross 1983). Presence of the only other species, A. carbonaria, in the plate-bound basins of Belgium suggests a northern hemisphere position for the Coyote Butte Formation and general affinity with the Carbonifer- ous—Permian Boreal fauna. Biogeographic evidence from so small a sample size is ten- uous, but corroborates preliminary paleo- magnetic evidence (Harbert et al. 1995) of a northern hemisphere position for the Oregon Blue Mountains region. Long dis- tance transport of the Grindstone terrane is not required for this distribution. Acteonina permiana, new species, from the Coyote Butte Limestone, Crook County, Oregon. 1, Systematic Paleontology Family Actaeonidae d’Orbigny, 1842 Genus Acteonina d’Orbigny, 1850 type species Acteonina carbonaria (DeKoninck, 1843) Acteonina permiana, new species Figs. 1-3 Diagnosis.—Very small (less than 4 mm length), shell outline like modern genus, Conus, with turret-stepped spire, elongate aperture, heterostrophic larval shell and col- umella with plica. Description.—Small for genus; subcy- lindrical outline, superficially resembling Conus; flat whorl profile and rectangular sutures, producing a turreted spire; broad sutural ramp; narrow aperture, shallowly expanded at base; concave, plicate colu- mella; heterostrophic larval whorl (Fig. 2). Depository.—Four silicified specimens VOLUME 111, NUMBER 4 are housed in the collections of the Depart- ment of Paleobiology, National Museum of Natural History (USNM), Smithsonian In- stitution, Washington, D.C. Type specimens include: USNM 498736 (holotype), USNM 498737 (paratype). The two other speci- mens (USNM 498738, USNM 498739) are fragments of turreted spires that also have the sutural ramp and five whorls. Measurements.—USNM 498736 (holo- type, Figs. 1, 2): length—4.10 mm, width— 3.61 mm, whorl number—6. USNM 498737 (paratype, Fig. 3): length—2.15 mm, width—1.66 mm, whorl number—5.5. Discussion.—The other species of the genus, A. carbonaria, is about two times the length of A. permiana. Whorl counts are the same for both species so we are not comparing juveniles of the Permian with adults of the Carboniferous. The acid etched residues from the Coyote Butte Fm. contain species from many different phyla that have variable lengths from several mm to several cm, so we are not biased by a size-sorted sample. Acteonina carbonaria also has a prominent collar-like ridge at the base of the parietal wall, which is absent or not preserved in A. permiana. Location.—The fossil locality is in Crook County, Oregon, which may be found on the Twelvemile Reservoir, Oregon 7.5 min- ute U.S. Geological Survey quadrangle, 1981 edition. The UTM coordinate is 11TKU75826907. The geographic coordi- nates are 119°47'17"N, 97°44'10"W. Age.—Early Permian (Wolfcampian to Leonardian) ages for the Coyote Butte For- mation are suggested by: brachiopods (Cooper 1957), conodonts (Wardlaw et al. 1982), colonial rugose corals (Stevens & Rycerski 1983) and fusulinids (Skinner & Wilde 1966). Literature Cited Blome, C. D., & M. K. Nestell. 1991. Evolution of a Permo-Triassic sedimentary melange, Grind- stone terrane, east-central Oregon.—Geological Society of America Bulletin 103:1280-1296. Cooper, G. A. 1957. Permian brachiopods from central 797 Oregon.—Smithsonian Miscellaneous Collec- tions 134(12):1-79. Harbert, W., J. Hillhouse, & T. Vallier. 1995. Paleo- magnetism of the Permian Wallowa Terrane; implication for terrane migration and oroge- ny.—Journal of Geophysical Research 100(7): 12573-12588. Hengstenberg, C. A., R. A. Hanger, E. E. Strong, & J. M. Yarnell. 1997. Paleogeographic significance of arthropods and molluscs of the Early Perm- ian Coyote Butte Formation, central Oregon.— Geological Society of America, Abstracts With Programs 29(3):23. Jones, D. L. 1990. Synopsis of Late Palaeozoic and Mesozoic terrane accretion within the Cordil- lera of western North America.—Philosophical Transactions of the Royal Society of London, series A 331:479—486. Knight, J. B. 1932. The gastropods of the St. Louis, Missouri, Pennsylvanian outlier: IV. The Pseu- domelaniidae.—Journal of Paleontology 6:189— 202. . 1936. Notes on Paleozoic Gastropoda.—Jour- nal of Paleontology 10:520—534. . 1941. Paleozoic Gastropod Genotypes.—Geo- logical Society of America, Special Papers 32: 1-510. Kollmann, H. A., & E. L. Yochelson. 1976. Survey of Paleozoic gastropods belonging to the subclass Opisthobranchia.—Annals Naturhistorische Museum Wien 80:207—220. DeKoninck, L. G. 1843. Description des animaux fos- siles, qui se trouvent dans le terrain carbonifére de Belgique, Liége, 1-651. . 1881. Fauna du calcaire carbonifére de la Bel- gique, Gastéropodes.—Annales Musée royal d’ Histoire naturel, 6:1—170. Meek, FB. 1863. Remarks on the family Actaeonidae, with descriptions of some new genera and sub- genera.—American Journal of Science 35:89— 94. Miller, E. L., M. M. Miller, C. H. Stevens, J. E. Wright, & R. Madrid. 1992. Late Paleozoic paleogeo- graphic and tectonic evolution of the western U.S. Cordillera. Pp. 57-106 in B. C. Burchfiel, P. W. Lipman, and M. L. Zoback, eds., The Cor- dilleran Orogen: Conterminous U.S. The geol- ogy of North America, volume G-3. Geological Society of America, Boulder, Colorado, 724 pp. D’Orbigny, A. 1842. Paléontologie Francaise, Terrains cretacés 2, Gastéropodes. G. Masson, Paris, 1— 456. . 1850. Prodrome de paléontologie stratigraphie universelle des animaux mollusques et rayonnés faisant suite au cours élémentaire de paléonto- logie. V. Masson, Paris, 1-392. Ross, C. A., & J. R. PB. Ross. 1983. Late Paleozoic accrted terranes of western North America. Pp. 798 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 7-22 in C. H. Stevens, ed., Pre-Jurassic rocks lonial rugose corals in western North Ameri- in western North American suspect terranes. ca—aids in positioning of suspect terranes. Pp. SEPM, Pacific Section, Los Angeles, 141 pp. 23—36 in C. H. Stevens, ed., Pre-Jurassic rocks Skinner, J. W., & G. L. Wilde. 1966. Permian fusuli- in western North American suspect terranes. nids from Pacific Northwest and Alaska. Part 2. SEPM, Pacific Section, Los Angeles, 141 pp. Permian fusulinids from Suplee area, east-cen- Wardlaw, B. R., M. K. Nestell, & J. T. Dutro, Jr. 1982. tral Oregon.—University of Kansas Paleonto- Biostratigraphy and structural setting of the logical Contributions Paper 4:11—15. Permian Coyote Butte Formation of central Stevens, C. H., & B. A. Rycerski. 1983. Permian co- Oregon.—Geology 10:13-16. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):799-806. 1998. Two new species of Spionidae (Polychaeta) from Tahiti, French Polynesia Pat Hutchings, Patrick Frouin, and Christian Hily (PH) The Australian Museum, 6 College Street, Sydney NSW 2000 Australia; (PF) Centre ORSTOM de Brest, B.P. 70, F-29280 Plouzane, France; (CH) URA CNRS D 1513, Institute d’Etudes Marines, 6 avenue Le Gorgeu, F-29287 Brest cedex, France Abstract.—Two new species of Spionidae, Scolelepis (Scolelepis) melasma and S. (S.) dicha are described from the intertidal area of exposed Tahitian beaches. The species co-occur and are restricted to basaltic sandy beaches. Scolelepis (S.) melasma is characterized by the absence of an occipital ten- tacle, neuropodial unidentate, hooded hooks beginning on setiger 40—41, no- topodial hooded hooks absent and branchiae fused basally. Scolelepis (S.) dicha also lacks an occipital tentacle, with neuropodial bidentate, hooded hooks beginning from setiger 26—32 and with posterior neuropodia having almost all hooded hooks, notopodal hooks absent, and branchiae fused ba- sally. During an ecological investigation of the intertidal fauna of exposed beaches in Ta- hiti, two extremely abundant species of spionids were collected. The polychaete fauna of French Polynesia or any other re- gion of the South Pacific is poorly known and many of the species that have been de- scribed are incompletely known. While at- tempting to identify these spionids using the key of Blake & Kudenov (1978), it be- came apparent that they represented two new species, of the genus Scolelepis Blain- ville, 1828 and the subgenus Scolelepis sensu stricto as defined by Maciolek i987): The following abbreviations have been used in the text, AM, Australian Museum, Sydney; BMNH, The Natural History Mu- seum, London; MNHN, Museum Nationale D’Histoire Naturelle, Paris; LACM, Los Angeles County Museum, Los Angeles; USNM, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. Scolelepis (Scolelepis) melasma, new species Fig. 1A—G, Table 1 Material examined.—French Polynesia, Tahiti, La Fayette Beach, 17°30.238’S, 149°27.377'W, intertidal, basaltic sand, 27 March 1995. Holotype (AM W23103), Paratypes: 2 (BMNH ZB 1996.518), 2 (MNHN UE413), 1 (USNM 170033), 10 (AM W23104), 1 (LACM Poly 1831, 1 (AM W23107). Description.—Description based on ho- lotype except where stated. Alcohol pre- served material pale yellow, live material dark green in colour. Holotype posteriorly incomplete, 23.0 mm in length, 1.52 mm in width anteriorly, with about 92 segments. Paratype material varying in length from 20 to 30 mm and 1.52 to 3.04 mm in width anteriorly for 63 to 95 setigers, none of the type material is complete and the majority of the type material consists of ripe males and females. Elongate pointed prostomium (Fig. 1A), 800 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON _ gg a Fig. 1. Scolelepis (Scolelepis) melasma new species. Holotype (AM W23103). A. Dorsal view of anterior segments, B. Lateral view of parapodium of setiger 3, C. Lateral view of parapodium of setiger 20, D. Lateral view of parapodium of setiger 50, E. Lateral view of parapodium of setiger 85, E Notoseta from setiger 3, G. 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Occipital papillae absent. Eye spots located just anterior to point of inflation. Peristom- ium expanded laterally to form pair of lat- eral wings enveloping the prostomium. Ventral mouth with raised glandular mar- gins, pharynx not extended on holotype but partially everted on some type material. Palps detached on holotype but bases where previously attached clearly visible. Palps on paratypes extending to between setigers 10-12. Setiger 1 with both noto- and neuropo- dia. Notopodia small rounded lobe with small bundle of capillaries, neuropodia, bluntly triangular lobe with capillaries. Se- tiger 2, with larger bluntly triangular neu- ropodial lobe and capillary setae, notopo- dium with small, rounded acicular lobe and laterally elongated postsetal lobe and a dig- itiform branchia attached basally, and nu- merous long capillary setae. Neuropodial and notopodial postsetal lobes increasing in size on subsequent setigers, margins of these lobes becoming thinner and crenulate. Similarly, the length of branchiae increas- ing on subsequent setigers and branchiae present on all remaining setigers of holo- type. Anterior branchiae fused basally to notopodial lamellae. Fig. 1B-—E illustrate the relative proportions of the parapodial lobes along the body. All notopodial setae capillaries, with smooth tips and narrow wings (Fig. 1F), se- tiger 3 with about 20 capillaries and setiger 85 with about five capillaries. Notopodial hooded hooks absent. Neuropodial setae initially capillaries; setiger 3 with about 12 capillaries; from setiger 41, unidentate hooded hooks appear (Fig. 1G), 2-3 per fascicle; setiger 85 with two capillaries and three hooded hooks. Segments well demarcated dorsally, with raised intersegmental ridges present, venter smooth with mid ventral glandular stripe. Holotype a gravid female with numerous oocytes restricted to the parapodia and the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON area just adjacent to the parapodia, from about setiger 38 onwards (Fig. 1D), oocytes restricted to the lateral walls. Variation.—In some paratypes a slight bulbous swelling occurs immediately below apex of the prostomium. The setiger on which neuropodial hooded hooks begin varies from 40 to 41. Posterior fragments collected indicate that the pygidium is a rounded cushion. Some ripe females with oocytes developed from setiger 34 and con- tinuing to about setiger 85. Ripe males with spermatozoa also restricted to parapodial lobes from about setiger 35 to 90. Remarks.—Maciolek (1987) reviewed the genus Scolelepis and based upon the structure of the hooded hooks, divided the genus into two subgenera, Scolelepis and Parascolelepis. The subgenus Scolelepis is characterized by falcate hooded hooks with Q—2 small apical teeth and a straight shaft, and the subgenus Parascolelepis is charac- terized by multidentate hooded hooks above a large main fang and curved shaft. Based upon these characters this new spe- cies from Tahiti belongs in the subgenus Scolelepis. Maciolek (1987) tabulated the major characteristics of the 37 described species and three other described species having unidentate neuropodial hooded hooks, S. (S.) knightjonesi (de Silva, 1961 incorrectly quoted as 1965 by Maciolek 1987) de- scribed from Sri Lanka, S. (S.) lamellicinata Blake & Kudenov, 1978 from Australia and S. (S.) lefebvrei (Gravier, 1905) from Dji- bouti, Red Sea. However, of these, S. (S.) knightjonesi has an occipital papilla present that is lacking in S. (S.) melasma. The new species also has shorter palps than S. (S.) knightjonesi, which has palps extending to setiger 28 rather than setigers 10—12; in ad- dition, the branchiae occur only laterally (Fig. 1A) and do not meet mid-dorsally, whereas on S. (S.) knightjonesi the bran- chiae overlap mid-dorsally. The new spe- cies can be easily distinguished from S. (S.) lefebvrei and S. (S.) lamellicinata as both these species have both noto- and neuro- VOLUME 111, NUMBER 4 podial hooded hooks, whereas the new spe- cies has neuropodial hooks only present. Since Maciolek’s (1987) review several additional species of Scolelepis (Scolelepis) have been described and these are shown in Table 1, using the same characters as used by Maciolek, together with the two new species described here. Two of these re- cently described species of Scolelepis (Sco- lelepis) have unidentate neuropodial hood- ed hooks, S. (S.) branchia Imajima, 1992 and S. (S.) lingulata Imajima, 1992. The first of these species has accessory bran- chiae, which clearly distinguishes it from S. (S.) melasma, and the latter species has neu- ropodial hooded hooks from setiger 21, whereas the new species has them present from setiger 40—41. Habitat and ecology.—The species oc- curs on all basaltic beaches in Tahiti which are found on both the east and west coasts of the island, on exposed beaches that are not protected by a barrier reef. The species lives in non-permanent mucous tubes and occurs in densities of 700—1000 m7, in fine sandy sediments in the intertidal region of the beach; the tidal range is about 50 cm. All these beaches are fully exposed to oce- anic waters and the worms occur in the surf zone. For more detailed ecological infor- mation see Frouin et al. (1998). Etymology.—The specific name is a Greek word for “black’’, which refers to the black basaltic sediments in which this species occurs. Distribution.—French Polynesia, Tahiti. Scolelepis (Scolelepis) dicha, new species Fig. 2A—G, Tables 1 & 2 Material examined.—French Polynesia, Tahiti, La Fayette Beach, 17°30.238’S, 149°27.377'W, intertidal, basaltic sand, 27 March 1995. Holotype (AM W23105), Par- atypes: 1 (BMNH ZB 1996.519), 1 (MNHN UE 414), 1 (USNM_ 170034), 20 (AM W23106) and 1 (LACM Poly 1830). Description.—Description based on holo- type except where stated. Alcohol preserved 803 material pale pink with scattered diffuse brown pigmentation on anterior dorsum and venter, most marked on prostomium. Palps detached but present with distinct brown pig- mentation along one axis. Live material dark brown with blood vessels clearly visible around parapodia. Holotype complete, 15.2 mm long, 1.52 mm wide anteriorly, with about 70 segments. Paratype material varies from 15 to 20 mm long and 1.14 to 1.7 mm wide anteriorly for 70 to 75 setigers. Prostomium pointed, simple elongated cone (Fig. 2A), peristomial wings poorly de- veloped. Occipital papillae absent. Pharynx partially extruded, thin walled and faintly pigmented. Two pairs of eye spots arranged on either side of caruncle. Caruncle extend- ing posteriorly, slightly raised forming an elongate occipital papilla on mid-axis of car- uncle. On either side of caruncle scars in- dicating point of attachment of palps. On paratypes some of which have palps still at- tached, these extend to setiger 27. Setiger 1 poorly developed, notopodia with small, rounded triangular lobe, noto- setae absent, neuropodia with small conical lobe with few capillary setae pesent, bran- chiae absent. Setiger 2 with elongate rounded notopo- dial lobe, and an elongate branchial lamella basally attached to notopodial lobe. Capil- lary notosetae numerous, forming a fan over the lobe. Neuropodia with semi-cir- cular lobe and neurosetal capillaries ar- ranged in a fan. Presetal lobes absent, both parapodial lobes postsetal in origin. Para- podial lobes and branchiae increasing in size on subsequent setigers (Fig. 2B), with margins of lobes thinner and margins of branchiae crenulated. Setiger 3 with about 12 notosetae and 10 neurosetae. Lobes and branchiae fully developed by setigers 3—4, branchiae not reaching mid dorsal line. Pos- teriorly, notopodial lobe becoming bilobed (Fig. 2C) but, by setiger 48 notopodial lobe almost entire again. Subsequently, the size of the parapodial lobes and branchiae de- clining although branchiae present almost to pygidium. 804 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cc Fig. 2. Scolelepis (Scolelepis) dicha new species. Holotype (AM W23105). A. Dorsal view of anterior segments, B. Lateral view of parapodium of setiger 3, C. Lateral view of parapodium of setiger 21, D. Lateral view of parapodium of setiger 48, E. Lateral view of parapodium of setiger 85, E Notoseta from setiger 3, G. Neuroseta from setiger 45, G. Dorsal view of pygidium. VOLUME 111, NUMBER 4 Table 2.—The relationship between body length, maximum anterior width and the setiger on which neu- ropodial hooded hooks are first present on S. (S.) di- cha. Start of Length Width Setigers hooks (set (mm) (mm) present no.) Comments 8.33 1.31 31* 31 gravid 8.33 1.31 34 29 11.90 0.60 SIl* 26 11.90 1.43 44* 31 13.10 1.43 42* 31 14.88 1.19 38* 29 15.48 1.19 64 29 17.50 1.31 70 30 18.45 1.31 70 30 19.05 1.55 55)" 32 gravid 19.64 1.31 69 31 gravid 20.24 1.90 47* 32 21.43 1.67 71 31 gravid 21.43 1.43 63 30 22.02 .5)5) oll 32 gravid 23.81 1.19 71 31 25.60 1.31 Vs 31 26.19 1.43 76 31 gravid 27.98 1.67 71 30 47.62 2.38 71 31 No. of individuals = 20. * Indicates incomplete individual. Notosetae capillaries throughout (Fig. 2B), broad bladed, narrow winged smooth tipped, with granular cores. Notosetal hooded hooks absent. Neurosetae initially capillaries re- placed with bidentate hooded hooks (Fig. 2F), which first occur on setiger 31, with a mixture of capillaries and hooks, but rapidly thereafter majority of neurosetae hooded hooks, about 5—6 per parapodium (Fig. 2D). Large part of dorsum uncovered with well demarcated, central longitudinal, dor- sal glandular stripe. Segments are relatively poorly defined, suggesting that internal sep- ta are not well developed. Venter smooth and internal body musculature clearly visi- ble through body wall. Holotype a gravid female with oocytes clearly visible from setiger 25, about 6—9 oocytes per setiger, but lacking from pos- terior segments. Pygidium small rounded cushion with two patches of eye spots (Fig. 2G). 805 Variation.—The setiger on which neuro- podial hooded hooks begin varies from 26 to 32 (Table 2) but on most individuals the hooks begin on setiger 29 or later. There is some evidence that with increasing size as measured by length, that the hooks begin lat- er, suggesting some setal replacement with in- creasing length and presumably age. As many of the individuals were posteriorly in- complete, the maximum width anteriorly was also measured, but this showed less support for the relationship between increasing size as measured by width and the later develop- ment of neuropodial hooded hooks. Based on the 11 complete individuals which were mea- sured, there was not a direct relationship be- tween total length of animal and maximum anterior width. This may be real or perhaps a function of preservation techniques em- ployed as in many polychaete species, the maximum anterior width can be used as a substitute for total length. Additional material would need to be examined to determine which was the valid interpretation. The intensity of the pigmentation pat- terns varies within the paratype material. In a ripe male the gametes are clearly visible from setiger 35 and are then present almost until the pygidium. Remarks.—As discussed above the genus Scolelepis has recently been reviewed by Maciolek (1987) and she erected two sub- genera. Scolelepis (S.) dicha belongs to the subgenus Scolelepis. Based on the data pre- sented by Maciolek (1987) and in Table 1, the following species in this subgenus pos- sess bidentate neuropodial hooded hooks, notopodial hooded hooks absent, occipital papillae absent and notosetae absent on se- tiger 1, thus resembling S. (S.) dicha: S. (S.) cantabra Rioja, 1918, S. (S.) vexillatus Hutchings & Rainer, 1979, S. (S.) variegata Imajima, 1992, S. (S.) williami de Silva, 1961. Pettibone (1963) examined the type of S. (S.) cantabra and found that the oc- cipital tentacle was lacking although Rioja had originally described it as being present, but she indicates that the holotype is dam- aged. However, regardless of this character, 806 the setiger on which neuropodial hooks Start is from setiger 20—25, whereas they start from setiger 26—32 in S. (S.) dicha. The new species can be separated from S. (S.) vexillatus, as the branchiae are always just fused basally, whereas in the latter spe- cies they are completely fused initially, only becoming partially free posteriorly from setiger 29. Scoloplos (S.) williami dif- fers from the new species as the neuropo- dial hooded hooks begin on setiger 40, whereas in the new species they begin from setiger 26—32. Finally, S. (S.) dicha can be separated from S. (S.) variegata by the de- gree of fusion of the branchiae, in the new species they are only fused basally, whereas in S. (S.) variegata they are completely fused with the notopodial lamellae until se- tiger 29. Thus S. (S.) dicha can be easily distinguished from all these species on the basis of the setigers on which neuropodial hooded hooks begin and the degree of fu- sion of the branchiae and is therefore de- scribed as a new species. Habitat and ecology.—The species oc- curs on the basaltic La Fayette beach in Ta- hiti and appears to be absent from other ba- saltic beaches that are found on both the east and west coasts of the island. The spe- cies lives in non-permanent mucous tubes and occurs in densities of 700-1000 m7’, in the upper intertidal region of the beach just below a line of rocks in fine sandy sedi- ments; the tidal range is 50 cm. This beach is fully exposed to oceanic waters and is not protected by an offshore barrier reef. The species co-occurs with Scolelepis (S.) me- lasma. For more detailed ecological infor- mation see Frouin et al. (1998). Etymology.—The specific name is a Greek word for “‘two”’ referring to the bi- dentate hooded hooks present in this species. Distribution.—French Polynesia, Tahiti, La Fayette Beach. Acknowledgments We should like to thank Anna Murray for illustrating the anterior views of both species and Kate Attwood for preparing the plates. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Blainville, H. de. 1828. Dictionnaire des Sciences Na- turelles 47:368—501. Blake, J., & J. Kudenov. 1978. The Spionidae (Poly- chaeta) from southeastern Australia and adja- cent areas with a revision of the genera.—Mem- oirs of the National Museum of Victoria. 39: 171-280. De Silva, P H. D. 1961. Notes on some polychaetes from Ceylon.—Spoila Zeylanica 29:164—194. Frouin, P., C. Hily, & P. Hutchings. (1998). Ecology of spionid polychaetes in the swash zone of ex- posed beaches in Tahiti (French Polynesia).— Comptes Rendus de |’Academie des Sciences de la vie. Paris 321:47—54. Gravier, C. 1905. Sur les annélides polychétes de la Mer Rouge (Cirratuliens, Spionidiens, Ari- ciens.—Bulletin du Muséum d’Histoire Natu- relle Paris 11:42—46. Hartmann-Schroéder, G. 1983. Die Polychaeten des Eu- litorals. Pp. 123-167 in G. Hartmann-Schréder & G. Hartmann, eds., Zur Kenntnis des Euli- torals der australischen ktisten unter besonderer Beriicksichtigung der Polychaeten und Ostra- coden. Teil 9. Die Polychaeten der antiborealen Sudkiiste (zwischen Albany im Westen und Ce- duna im Osten.—Mitteilungen aus dem Ham- burgischen zoologischen Museum und Institut. 80. . 1991. Beitrage zur Polychaetenfauna der Ba- hia Quillaipe (Siid-Chile)—Helgolander Meer- esuntersuchungen 45:39—58. Hutchings, P. A., & S. F Rainer. 1979. The polychaete fauna of Careel Bay, Pittwater, New South Wales, Australia——Journal of Natural History 13:745-779. Imajima, M. 1992. Spionidae (Annelida, Polychaeta) from Japan VIII. The genus Scolelepis.—Bul- letin of the National Science Museum. Series A (Zoology). 18(1):1—34. Maciolek, N. J. 1987. A redescription and records of Scolelepis (Polychaeta: Spionidae) from east coast of North America, with a review of the subgenera. Pp. 16—40 in K. Fauchald, ed., Bul- letin of the Biological Society of Washington No. 7. Pettibone, M. H. 1963. Revision of some genera of polychaete worms of the family Spionidae, in- cluding the descriptions of a new species of Scolelepis.—Proceedings of the Biological So- ciety of Washington 76:89—104. Sikorski, A. V. 1994. New arctic species of Scolelepis (Polychaeta, Spionidae). Pp. 123-167 in J.-C. Dauvin, L. Laubier, & D. J. Reish, eds., Actes de la 4¢mes Conférence internationale des Po- lychétes—Mémoires Museum Nationale d’ Histoire Naturelle 162. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):807—822. 1998. New species of bathyal and abyssal Sabellariidae (Annelida: Polychaeta) from near New Caledonia (southwest Pacific Ocean) Jean-Paul Lechapt and David W. Kirtley+ (JPL) Laboratoire maritime de Dinard, Museum National d’ Histoire Naturelle, 17 Avenue George V, F-35800 Dinard, France; (DWK) PO. Box 2713, Vero Beach, Florida 32961-2713, U.S.A. Abstract.—Specimens described in this work were collected during the BIO- CAL and BIOGEOCAL cruises (Biology and Geology of New Caledonia) in 1985 and 1987 conducted by the Museum National d’ Histoire Naturelle (Paris, France). All of the 488 specimens collected from depths between 440 m and 1870 m belong to the subfamily Lygdaminae Kirtley, 1994, which is charac- terized by the presence of four parathoracic segments. Among these specimens, we recognize four genera, including Bathysabellaria Lechapt & Gruet, 1993 with two species, a new species of Lygdamis Kinberg, 1867, two new species of Phalacrostemma Marenzeller, 1895, and a new species of Tetreres Caullery, 1913. All of these species are set apart from other known species by the shape of the outer and inner paleae and by their geographic and bathymetric distri- butions. These collections illustrate the uniqueness and richness of the poly- chaete fauna of the Sabellariidae of this Pacific area. Except for the relatively limited accounts of Hoagland (1920:627), Treadwell (1926: 191), Caullery (1944:54—66) from the Phil- ippines and Indonesia, and that of Gibbs (1971:101—211) from the Solomon Islands, the benthic polychaete fauna from great depths of this geographic area remains poorly known, especially for the family Sabellari- idae. The material on which this study is based was collected during the 1985 and 1987 cruises of the “Biology of New Cale- donia” project, BIOCAL and BIOGEOCAL, respectively, conducted by the French Na- tional Museum of Natural History of Paris. Specimens were collected with a Waren- type rock dredge (Stations DW) or with a beam trawl (Stations CP), fixed in a sea- water formalin solution and stored in 70% ethanol. The material was received from the Centre National de tri d’Oceanographie 7 Deceased July 1997. Biologique, Brest, France, examined under low power magnification and selected mor- phological features were studied by scan- ning electron microscopy (SEM). All type material is deposited in the National Mu- seum of Natural History, Paris (MNHNP). Family Sabellariidae Subfamily Lygdaminae Kirtley, 1994 The family Sabellariidae was subdivided by Kirtley (1994:14) into two subfamilies, the Sabellariinae with three parathoracic segments and the Lygdaminae with four parathoracic segments. All specimens con- sidered here belong to the Lygdaminae, the genera being distinguished as follows: la. Compound (branched) feeding tentacles oF ean tag a i8h Sila del ugk eter a nate Lygdamis 1b. Simple (unbranched) feeding tentacles 2) 2a. Operculum completely divided i GAR Aas Sit lsc deb 3 Phalacrostemma 2b. Operculum not completely divided ... 3 808 NS \ \ * Fig. 1. et! PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lygdamis splendidus, new species, (Holotype, MNHNP UC 356): A, ventral view of anterior end; B, dorsal view of anterior end; C, left lateral view; D, surface view of crown; E, cross-sectional diagram showing position of paleae and nuchal hooks on crown. Scale bars = 1 mm. Abbreviations: ip = inner paleae, mo = median organ, nh = nuchal hook, op = outer paleae, ps = parathoracic setiger. 3a. Operculum completely fused along ventral margin Bathysabellaria Operculum with deep indentation along ventral margin Tetreres S10). Genus Lygdamis Kinberg, 1867 Lygdamis splendidus, new species Ricsal, 2 Material examined.—Southwest off New Caledonia, BIOCAL cruise, station DW 66, 24°55'S, 168°21’E, 515 m, 3 Sep 1985, Ho- lotype (MNHNP UC 356), Paratypes (MNHNP UC 357) (two specimens). Description.—Holotype incomplete, missing posterior most segments and cau- da; total length 13 mm, thoracic diameter 3 mm. Anterior end of opercular stalk completely divided into bilaterally sym- metrical lobes (Fig. 1A, B). Opercular pa- leae of two kinds, in curved, dorsoventral rows (Fig. 1C, D). Outer paleae, 100—120 in number, 2 mm long, transparent, smooth, with distal ends terminating in fine tips (Fig. 2A, C); inner paleae, 36—40 in number, 3 mm long, yellow-gold, smooth, with slightly curved tips (Fig. 2B, D). Falcate dorsal nuchal hooks, tips re- curved inward toward dorsal midline (Fig. 1E). With well-developed median organ arising from dorsal sagittal suture between opercular lobes (Fig. 1B). Twenty-five pairs of compound feeding tentacles on ventral margins of each side of opercular peduncle, pair of short prehensile prosto- mial tentacles arising from anterior margin of upper lip of stoma. Large U-shaped building organ with conical lobes on lat- VOLUME 111, NUMBER 4 809 Fig. 2. Lygdamis splendidus, new species, (Holotype, MNHNP UC 356): A, outer paleae; B, inner paleae; C, SEM of outer paleae; D, SEM of inner paleae. Scale bars = 0.1 mm for A, B and 100 pm for C, D. 810 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON eral margin, with conical cirrus with small bundle of short, ventral, finely serrate cap- illary setae. Setiger 2 with same bundle of finely serrate capillary setae. Four para- thoracic segments (Fig. 1C) with small cir- ri on distal dorsal margin of notopodial sheaves; 7—8 paired, stout setae with lan- ceolate distal tips and spinous, capillary companion setae with curved tips in trans- verse row on each notopodial sheave. Neu- ropodial setae similar, but smaller in size and fewer in number. Dorsal branchiae on parathoracic segments. Abdominal seg- ments with bundles of long, spinous neu- rosetae and uncinigerous notopodial tori. Cauda not observed. No tubes. Etymology.—The specific name, from Latin splendidus = magnificent, refers to its very beautiful appearance. Remarks.—Lygdamis splendidus is clear- ly distinguished from other species of Lyg- damis by the shape of the outer paleae, ta- pering to fine distal tips, the shape of inner paleae, and total number of paleae; more- over, of all Lygdamis species, L. splendidus occurs at the greatest depth. L. indicus Kin- berg, 1867 and L. gilchristi (McIntosh, 1922) appear to be most similar to L. splen- didus but differ in the shapes of the inner and outer paleae and by their geographical and bathymetric distributions (Fig. 3). Key to the species of Lygdamis [after Kirtley (1994:117—-118)] la. Outer paleae with smooth lateral mar- AIS Pie cur Sop eee ene ate keh eee 4 1b. Outer paleae with lateral margins not STO OUE Ey kh ea aks ccs ei Ala Soa 2 2a. Outer paleae with serrate lateral mar- gins, with slightly concave inner sur- faces (A, Fig. 3) (West Africa, 77 m) Bee ol A Grae Eee perthes cena ee L. robinsi 2b. Outer paleae with lateral margins not SEihAlGy Si, Purwhetcee vse ee line ee 3 3a. Outer paleae with complex ornamen- tation, with irregular, wavy, transverse lines across middle portion of blade, inner paleae with fine, marginal dec- oration (B, Fig. 3) (West Africa, 22 m) L. kirkegaardi 3b. Without complex ornamentation on outer paleae 4a. Distal ends of outer paleae straight, nearly bilaterally symmetrical ...... > 4b. Distal ends of outer paleae curved, conspicuously asymmetrical 5a. Outer paleae with narrow distal spines terminating in narrow, delicate spike; nuchal hooks with weakly recurved tips (C, Fig. 3) (Indonesia, 36 m) ... shea) nae eh o's chads eee L. indicus 5b. Outer paleae with distal tips tapering to very long spike; nuchal hooks with strongly recurved tips; inner paleae with slightly curved tips (D, Fig. 3) (NewaGaledoniay 55m) ieee 5 15 EE TEE L. splendidus, new species 5c. Outer paleae converge slowly to sharp tips; inner paleae converging abruptly to symmetrical short, sharp tip (E, Fig. 3) (South Africa, 75 m).... L. gilchristi 6a. Outer paleae with blade margins al- most parallel through middle portion, terminating in narrow mucronate tips; inner paleae with gently bent tips (E Fig. 3) (Australia, 70 m) ... ZL. augeneri 6b. Outer paleae narrowing continuously from middle portion to tips ........ 7 7a. Distal ends of outer paleae slightly asymmetrical; inner paleae with bent distal ends; nuchal hooks strongly re- curved, with sharp tips (G, Fig. 3) (Australia, 64 m) L. giardi 7b. Outer paleae moderately expanded through middle portion and tapering slowly to sharp distal point; inner pa- leae tapering slowly to dull point; nu- chal hooks with strongly recurved sharp tips (H, Fig. 3) (England) .... Oa BANS ae a ayes Wiha eeeeeeeemene L. muratus 8a. Outer paleae with distal tip as flat pro- jection from cylindrical paleal blade 9 8b. Outer paleae with distal tip as cylin- drical continuation of paleal blade .. 12 9a. Middle portion of outer paleae ex- panded, distal %4 constricted and slightly curving (I, Fig. 3) (Madagas- Cal). fn cane 1g eee ae L. malagasiensis 9b. Distal % of outer paleae not abruptly constricted “2 2 x. Day eer ee 10 Distal tip of outer paleae very narrow, rounded at distal end of blade, with elongate, sigmoidal spike; inner pale- 10a. VOLUME 111, NUMBER 4 Fig. 3. vations. A = L. robinsi, B = L. kirkegaardi, C = L. indicus, D = L. splendidus, new species, E = L. gilchristi, F = L. augeneri, G = L. giardi, H = L. muratus, | = L. malagasiensis, J = L. bhaudi, K = L. ehlersi, L = L. gibbsi, M = L. nesiotes, N = L. laevispinis, O = L. rayrobertsi, P = L. dayi, Q = L. curvatus. 10b. lla. 11b. 12a. 12b. 13a. ae with narrow tips (J, Fig. 3) (Mad- agascar) L. bhaudi Distal tip of outer paleae without sig- moidal spike Outer paleae with narrow, curved dis- tal tip (K, Fig. 3) (Indonesia, Philip- pines, 45 m) L. ehlersi Outer paleae with wide, curved distal tip; tip of nuchal hook bent about 90° from axis of shaft (L, Fig. 3) (Solo- mon Islands, coral reef) L. gibbsi Distal tips of outer paleae conical, sig- moidal in outline Distal tips of outer paleae not sigmoi- dal in outline Outer paleae with sigmoidal tip, curved inward about 30° from vertical 13b. 14a. 14b. 811 * Outer paleae ig Inner paleae. Shapes of outer paleae and inner paleae of species of the genus Lygdamis with geographical obser- axis of shaft toward center of crown; nuchal hooks very large, strongly bent to blunt tips (M, Fig. 3) (East Pacific Ocean, coral reef) L. nesiotes Outer paleae with sigmoidal tip, curved inward about 80° from vertical axis of shaft toward center of crown; nuchal hooks relatively small, strongly bent, short, stout, with sharp tip (N, Fig. 3) (South Central Pacific Ocean) L. laevispinis Tip of outer paleae with stout, short abruptly pointed tips; nuchal hooks elongate, large, strongly bent (O, Fig. 3) (Florida, Gulf of Mexico, 328 m) L. rayrobertsi Tips of outer paleae short, blunt .... 15 812 15a. Inner paleae with fine transverse mark- ings (P, Fig. 3) (South Africa) ... L. dayi 15b. Inner paleae stout, tapering to dull point; outer paleae with stout, rounded tips (Q, Fig. 3) (Bonin Islands) Genus Phalacrostemma Marenzeller, 1895 Phalacrostemma tenue, new species Figs. 4, 5 Material examined.—Southeast off New Caledonia, BIOCAL cruise, station CP 62, 24°19'S, 167°48’E, 1395 m—1410 m, 2 Sep 1985, Holotype (MNHNP UC 358), Para- types (MNHNP UC 359) (two specimens). Description.—Holotype 4 mm long; di- ameter of opercular crown 1.5 mm. Anterior end of opercular stalk completely divided into two bilaterally symmetrical halves (Fig. 4A, B). Each side of crown with 16—17 outer paleae; 2—3 inner paleae in dorsal position. Outer paleae with transverse striations and expanded distal margins, with regular distal denticles forming fringe (Figs. 4C, 5A, B). Inner paleae smooth, tips blunt with regu- larly spaced transverse ridges (Fig. 4D). Two nuchal hooks on each side of anterior dorsal midline of stalk, each curving to an acute tip. Robust median organ arising from be- tween dorsal inner margins of stalk. Pair of long prehensile prostomial tentacles arising from anterior margin of upper lip of stoma, with 4 simple, ciliated feeding tentacles on each side of buccal cavity. First setiger with bundle of ventral pecti- nate capillary setae. Second setiger with conical cirrus with bundles of pectinate se- tae. Four parathoracic segments, each with neuropodial sheaves of setae with pectinate distal margins; notopodial sheaves with 5—7 setae with oar-shaped denticulate and acute distal tips, with companion capillary setae. Eight abdominal setigers with bundles of long, slender and spinous capillary neuro- setae, and uncinigerous notopodial tori, un- cini with 7—8 teeth in double row. Lacking cauda. No tubes. Paratypes 4 mm long with three abdomi- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nal setigers. Only two inner paleae on right side of stalk, 18 outer paleae on left side of stalk. Etymology.—The specific name, from Latin tenuis = thin, refers to its very small size. Remarks.—Phalacrostemma tenue 1s closely related to P. lechapti Kirtley, 1994, formerly described as P. cidariophilum Fau- vel, 1914 (not Marenzeller, 1895), but the arrangement of spinous fringes on the outer paleae and the parathoracic notosetae (Table 1) clearly distinguish these two abyssal forms, one from the Atlantic (Fauvel 1914: 273), the other from the Pacific. Phalacrostemma profundum, new species Fig. 6 Material examined.—Loyauté Basin, zone Z2, BIOGEOCAL cruise, station CP 317, 20°48'S, 166°53’E, 1630 m, 2 May 1987 (one specimen); off southwest Lifou, zone SB2, BIOGEOCAL cruise, station CP 265, 21°04'S, 167°04’E, 1870 m, 18 Apr 1987 (two specimens with tubes), Holotype (MNHNP UC 360), Paratype (MNHNP UC 362); off southwest New Caledonia, zone SB5, BIOGEOCAL cruise, station CP 214, 22°44'S, 166°28’E, 1665 m, 9 Apr 1987 (one specimen), Paratype (MNHNP UC 361). Description.—Holotype 12 mm long, in- cluding opercular crown; diameter of oper- cular crown 2 mm, thoracic diameter 1.8 mm. Short opercular stalks well divided into two symmetrical halves with outer paleae 2 mm long (Fig. 6A, B). Each half of crown with 30—40 outer paleae spirally arranged, some paleae moved backward in relation to external row. These outer paleae tapered, ringed, without spinous ornamentation (Fig. 6D, E). Along dorsal edge of each crown, 2-3 yellow, smooth, inner paleae with distal apex slightly curved (Fig. 6C). Crown mar- gins with 13-14 long bifid pericoronal palpi. Four pairs of curved dorsal hooks. A small median organ present between dorsal inner margins of stalk. Pair of large, deeply VOLUME 111, NUMBER 4 813 Fig. 4. Phalacrostemma tenue, new species, (Holotype, MNHNP UC 358): A, ventral view of anterior end; B, dorsal view of anterior end; C, surface view of crown; D, inner paleae. Scale bars = 1 mm for A—C and 0.1 mm for D. See Fig. 1 for key to abbreviations. grooved, peristomial tentacles arising from anterior margin of stoma. Four feeding ten- tacles on each anterior lateral margin of buc- cal cavity. Large glandular, U-shaped build- ing organ with marginal bundles of short spinous capillary setae. Setiger 2 with bun- dles of spinous capillary setae and short tri- angular cirri. $14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Phalacrostemma tenue, new species, (Holotype, MNHNP UC 358): A, SEM of outer paleae; B, SEM of outer paleae magnified. Scale bars = 100 ym for A and 10 pm for B. Four parathoracic segments with slightly spinous neuropodial capillary setae and bun- dles of 6—7 tapering bristly notopodial setae with smooth capillary companion setae. Ab- dominal segments with spinous capillary neurosetae, notopodia with comblike uncini with 8—9 rows of teeth. Cauda absent. Tubes composed of sand grains. Etymology.—The specific name, from the Latin profundus = deep, refers to the great depths from which it occurs (1720 m on av- erage). Remarks.—Phalacrostemma profundum, new species, is distinguished from P. ci- dariophilum Marenzeller, 1895 and from other species of this genus by the smooth outer paleae and the shape of the inner pa- leae and by its geographical distribution (Table 1). Genus Bathysabellaria Lechapt & Gruet, 1993 Characteristics of this genus include an opercular stalk completely fused; blades of outer paleae broad, stout, inner surfaces slightly concave, terminating in acuminate tips, blades with basal transverse ridge, ro- tated around longitudinal axis of shafts; in- ner paleae twice as long as outer paleae, in- ner surface slightly concave, outer surfaces expanded, gently curved toward center of crown, with 3—4 simple, small, filamentous ciliated feeding tentacles on each side of buccal cavity; two relatively small preoral tentacles; nuchal spines straight, cylindrical, with very slightly curved distal tips; four parathoracic segments (Lechapt & Gruet 1)33))\. Bathysabellaria neocaledoniensis Lechapt & Gruet, 1993 Fig. 7 Material examined.—Southwest Pacific Ocean: off New Caledonia, BIOCAL cruise, stations DW 44, 22°47'S, 167°14’E, 440 m—450 m, 30 Aug 1985, 24 specimens, DW 77, 22°15'S, 167°15’E, 440 m, 3 Sep 1985, one specimen. Description.—Opercular paleae of two kinds: outer paleae in concentric row along anterior periphery of crown surface, 28 in number, bright yellow-gold, smoothly ta- VOLUME 111, NUMBER 4 Table 1—Some characteristics of species of Phalacrostemma Marenzeller, 1895. 815 SPECIES Im fla ila] i la] Ta] Tla:) ila] Io a=] ‘la-) ila) tla=} abyssalis . cidariophilum . dorothyae . gloriae - gwendolynae . lechapti » paulinae - perkinsi . setosa . tenera - profundum . tenue Kirtley, 1994 Marenzeller, 1895 Kirtley, 1994 Kirtley, 1994 Kirtley, 1994 Kirtley, 1994 Kirtley, 1994 Kirtley, 1994 (Treadwell, 1906) (Augener, 1906) this work this work Characteristics of outer paleae With fringe with simple short distal omamentation Smooth distal margins slightly inclined from horizontal Thecae of middle part of blade with short finely denticulate fringe forming crescentic distal margin Thecae of middle portion with shorter and fewer denticles slightly undulatory horizontal with trace Distal fringe with expanded delicately attenuate distal extension Distal of outer paleae with irregularly distally expanded rows of. alternately wide and narrow marginal extensions, with both long and short, pectinate, distal tips Distal portion with expanded distal regular denticles forming distal fringes Middle portion without distinct dentition and very weakly expanded distal margins Distal portion without conspicuously expanded, with distal margins forming horizontal zig-zag pattern. Incomplete description Tapering, ringed without spinous omamentaion Longitudinal striations and expanded distal margins with regular distal denticles forming fringes Aspect of outer 20 pm paleae Hy 20 um 10pm Number of outer paleae no data no data no data no data no data no data no data Characteristics of inner paleae no data Flattened, with gently rounded straight tips Irregular, slightly inflated thecae with distal margins Thecae well developed with distal margins producing zig-zag pattern no data With conspicuous thecae and with deep groove along proximal median portion no data no data no data no data Smooth with distal portion lightly incurved Smooth blunted tips with regularly spaced transverse markings Geographic occurence Flores Sea (Indonesia) Adnatic, Mediterranea Horida Gulf of Mexico Carmbean Sea Azores West Atlantic Bahamas Islands Hawai Islands Barbados Off New Caledonia Off New Caledonia Depth sampling 794m 485 m—1298 m 230m 228m 880 m—1440 m 2160 m 1360 m 300 m—700 m 1700 m 1400 m 816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Phalacrostemma profundum, new species, (Holotype, MNHNP UC 360): A, ventral view of anterior end; B, dorsal view of anterior end; C, surface view of crown; D, SEM of outer paleae magnified; E, SEM of outer paleae. Scale bars = 1 mm for A—C, 10 wm for D and 100 pm for E. See Fig. 1 for key to abbreviations. VOLUME 111, NUMBER 4 Qa St See 817 Fig. 7. Bathysabellaria neocaledoniensis: A, ventral view; B, left lateral view; C, dorsal view of anterior end; D, view of anterior surface of crown; E, cross-section of anterior end of crown showing position of opercular setae and nuchal spines. Scale bar = 2 mm. See Fig. 1 for key to abbreviations. pered distal margin, and frayed or broken tips; inner paleae in single irregularly crowded row, 11 in number, erect, twice as long as outer paleae (Fig. 7A—E); blades in- flated, terminating in slightly bent, mucro- nate tips, with basal concavity and numer- ous fine transverse serrations, with simple pair of golden, acicular nuchal spines on each side of dorsal midline of opercular stalk. 818 ~ rag? aN og Ss \\: BB / iil j f (0 eae IP Sites Fig. 8. = 1 mm. Agglutinated tubes composed of small sand grains and Globigerina tests. Remarks.—Known only from the locali- ties reported herein. Bathysabellaria spinifera Lechapt & Kirtley, 1996 Fig. 8 Material examined.—Off New Caledo- nia, BIOCAL cruise, stations DW 36, 23°08'S, 167°11'’E, 650 m—680 m, 29 Aug 1985; 11 specimens;) D W533; 23 1008S. 167°10’E, 675 m—680 m, 29 Aug 1985, 3 specimens; DW 46, 22°53’S, 167°17'E, 570 m—610 m, 30 Aug 1985, 145 specimens; DW 51, 23°05’S, 167°45’E, 680 m—700 m, 30 Aug 1985, 287 specimens and MUSOR- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bathysabellaria spinifera: A, ventral view of anterior end; B, dorsal view of anterior end. Scale bars STOM IV cruise (N/O Vauban), station DW 222, 22°57'S, 167°33’E, 410 m—440 m, 30 Sep 1985, 2 specimens. Description.—Opercular crown with two kinds of paleae: outer paleae bright yellow, blades smoothly tapered, with fine serra- tion, 33—51 in number in various specimens and disposed in a complete circle around periphery of crown without any interruption or ventral indentation (Fig. 8A, B); inner paleae golden brown, arranged in two or three concentric rings, not in a spiral whorl; blades spiniform, distally curved, with in- flated base and numerous fine serrations, one or two short inflated paleae inserted be- tween long curved paleae, 20—30 in exter- nal row, 10 to 20 in the internal row. Two VOLUME 111, NUMBER 4 819 Cc Fig. 9. Tetreres robustus, new species, (Holotype, MNHNP UC 363): A, view of surface of crown; B, ventral view; C, cross-section of anterior end of crown showing position of opercular setae and nuchal spines. Scale bars = 1 mm for A, C and 10 mm for B. Straight, golden, cylindrical, acicular nuchal Remarks.—Bathysabellaria spinifera, spines with slightly curved distal tips on only from the localities reported herein (Le- each side of the dorsum of the opercular chapt & Kirtley 1996), is characterized by stalk. a greater number of individuals (400 spec- No tubes were sampled. imens) and a wider bathymetric distribution 820 than the other species of the genus, B. neo- caledoniensis. Genus Tetreres Caullery, 1913 Tetreres robustus, new species Figs. 9-10 Material examined.—Southeast off New Caledonia, BIOCAL cruise, stations DW 44, 22°47'S, 167°14’E, 440 m—450 m, 30 Aug 1985, 2 specimens (Holotype MNHNP UC 363); DW 77, 22°15'S, 167°15’E, 440 m, 5 Sep 1985, 5 specimens (Paratypes MNHNP UC 365); blocks with burrows in- side (UC 364). Description.—Holotype 35 mm long with 19 setigers (incomplete specimen). Thoracic diameter 8 mm, opercular crown 8 mm in diameter. Opercular stalk partially divided along dorsal midline (Fig. 9A). Outer paleae, 70—80 in external series, yel- low gold, smooth, with distally spatulate blades (Figs. 9A, B, 10A, B); inner paleae, 3—4 on each side, parallel to midline of crown, 6 mm long (Figs. 9A, 10D). Pair of large dorsal nuchal hooks bent outward at right angles to longitudinal axis of blades and shaft with tips pointing inward toward prostomium (Figs. 9C, 10C). Opercular stalk with 25—30 long, conical papilli; relatively short peristomial tenta- cles, 12 in number, and 2 long paired lateral lips. Building organ with accessory setig- erous cirri on each side. Series of 4 elon- gate, triangular cirri along either side of se- tiger 2; inferior cirri with tiny bundles of fine, spinous capillary setae. Four parathor- acic segments with fin-like notopodia, each with 7—9 stout setae with lanceolate tips and companion capillary setae. Neuropodia with similar setae, fewer in number (4—6), with pair of dorsal branchiae on each para- thoracic segment. Abdominal segments with neuropodial cirri with bundles of long capillary setae, some long, smooth capillary setae, and strongly spinous capillary setae. Notopodial uncini with 8—9 teeth in double rows. All specimens were collected inside bur- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rows of 6—7 mm in diameter, in large blocks made of Globigerina tests (UC 364). Etymology.—The specific name, from the Latin robustus = robust, refers to its great size. Remarks.—Tetreres robustus, new spe- cies, is characterized by the number and shape of the outer paleae and belongs to the group of species whose outer paleae pos- sess elongate, flattened blades. Key to the species of Tetreres la. Outer paleae with elongate flattened blades :-. 0... oat oe Z 1b. Outer paleae without flattened blades 4 2a. Nuchal hooks with short distal tips (Indian Ocean, 896 m) . T. sandraae 2b. Nuchal hooks with long distal tips .. 3 3a. Blades of outer paleae with attenuate distal spike. ...5.... 09:2 oe 8 3b. Blades of outer paleae without atten- uate. distal ‘spike .< =... -- ae 6 4a. Cylindrical outer paleae with distal prolongation °? 2.22. 25 eee 5 4b. Cylindrical outer paleae without distal prolongation (Indonesia, 330 m) .... wat oe & babs ia be ele eee T. superbus 5a. Distal tip of outer paleae short, flat- tened, with filiform plume (Philippine Islands; 1470 m) 7225552 T. philippinensis 5b. Blades of outer paleae short, flattened, without filiform distal tip (Antarctica, 3803.m). ... .. die ae T. maryriceae 6a. Blades of outer paleae taper abruptly to shghdy flattened tip =-3- eee Ji 6b. Blades of outer paleae taper gradually to acute tip?. 22: 2. 2... eee 8 7a. Blades of inner paleae with internal fusiform outline (Western Atlantic, AS25 yt ae eee T. varians 7b. Blades of inner paleae without internal fusiform outline (Hawaiian Islands, SSOuml) eee ss ers eee eee T. baileyae 8a. Margins of blades of outer paleae al- most parallel 8b. Margins of blades of outer paleae not parallel 9a. Outer paleae with acute extended tip (Antarctica, 4758 mm). ~.- -. aan T. cassidyi 9b. Outer paleae with dyssimetrical acute tip (Southeast Pacific Ocean, 440 m) T. robustus, new species VOLUME 111, NUMBER 4 821 Fig. 10. Tetreres robustus, new species, (Holotype, MNHNP UC 363): A, SEM of anterior end; B, SEM of outer paleae; C, SEM of surface of crown; D, SEM of inner paleae enlarged. Scale bars = 1 mm for A, C and 100 pm for B, D. See Fig. 1 for key to abbreviations. 10a. Margins of blades of outer paleae across surface (Northwest Atlantic wide through distal one-half (East Ocean; 4825 mM) sans see oe T. perryi Central Pacific Ocean, 3850 m) NPN Mite cats Tc Ro Oa nels T. jirkovi Acknowledgments 10b. Margins of blades of outer paleae wide through middle portion, with The authors are indebted to J. Le Lannic conspicuous transverse thecal bands from the Centre Commun de Microscopie 822 Electronique 4 Balayage de l’université de Rennes I. France, for assistance with the scanning electron microscopy. I would like to dedicate this work to the memory of Da- vid, who died during the preparation of this manuscript, and with whom I have worked on the Sabellariidae for several years. With- out him this work would never have been completed. Literature Cited Caullery, M. 1944. Polychétes sédentaires de VExpédition du Siboga: Ariciidae, Spionidae, Chaetopteridae, Chloremidae, Opheliidae, Ow- eniidae, Sabellariidae, Sternaspidae, Amphic- tenidae, Ampharetidae, Terebellidae.—Siboga- Expeditiie, Leiden 24(2):1—204. Fauvel, P. 1914. Annélides polychétes non pélagiques provenant des campagnes de |’ Hirondelle et de la Princesse Alice (1885—1910).—Résultats des Campagnes Scientifiques Monaco 46:1—432. Gibbs, P. E. 1971. The Polychaete worms of the Sol- omon Islands.—Bulletin British Museum (Nat- ural History), Zoology 1(5):101—211. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hoagland, R. A. 1920. Polychaetous annelids collected by the US Fisheries steamer Albatross during the Philippine Expedition of 1907 to 1910.— Bulletin of the United States National Museum 100(1):603—634. Kirtley, D. W. 1994. A review and taxonomic revision of the family Sabellariidae Johnston, 1865 (An- nelida; Polychaeta)—Vero Beach, Florida, Sa- becon Press Science Series, 223 pp. Lechapt, J. P, & Y. Gruet. 1993. Bathysabellaria neo- caledoniensis, a new genus and species of Sa- bellariidae (Annelida Polychaeta) from bathyal zones off New Caledonia (southwest Pacific Ocean).—Zoologica Scripta 22(3):243—247. , & D. W. Kirtley. 1996. Bathysabellaria spi- nifera (Polychaeta: Sabellariidae), a new spe- cies from deep water off New Caledonia, south- west Pacific Ocean.—Proceedings of the Bio- logical Society of Washington 109:560—574. Treadwell, A. L. 1926. Contributions to the Zoology of the Philippine Archipelago and adjacent re- gions. Additions to the polychaetous annelids collected by the US Fishery Steamer Albatross, 1907-1910, including one new genus and three new species.—Bulletin of the United States Na- tional Museum 100(6):183—193. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):823-828. 1998. Two new species of Nereis (Polychaeta: Nereididae) from Todos Santos Bay, Ensenada, Baja California, México J. A. de Le6n-Gonzalez and V. Diaz-Castanfeda (JAL-G) Division Biologia Marina, Centro de Investigaciones Bioldégicas del Noroeste, S.C. Apartado Postal 128, La Paz, Baja California Sur, 23000, México; (VD-C) Departamento de Ecologia Marina, Centro de Investigaciones Cientificas y de Educacion Superior de Ensenada, Apartado Postal 2732, Ensenada, B.C. 22860 México Abstract.—The nereidid fauna of Todos Santos Bay was studied. In this paper we describe two species of Nereis that are new to science. Nereis fau- chaldi belongs to the species group with the dorsal ligule expanded in posterior parapodia, and homogomph falcigers with smooth blades. Nereis imajimai be- longs to the species group with the dorsal ligule short in posterior parapodia, and homogomph falcigers with short and dentate blades. Nereididae is one of the polychaete fam- ilies with a broad distribution. Members of this family are found at all latitudes and from the intertidal to abyssal depths; how- ever, they are found more frequently in Shelf depths. Terminology of parapodial structures follows Hutchings & Reid (1990). Baja California, Mexico, is a state with two different coasts bordering two different water masses, the Gulf of California (Cortes Sea) on the eastern side, and the Pacific Ocean on the western side. Nereidid poly- chaetes have been little studied on the Pa- cific side. Only a few species have been re- ported by Hartman (1952, 1963), Treadwell (1923), Berkeley & Berkeley (1958) and Reish (1963). In the present work we de- scribe two new species of Nereis from To- dos Santos Bay, located in the Pacific coast of Baja California, 100 km south of the U.S.A.-Mexico border. The study is based on samples from 47 stations collected be- tween 31°40’ to 31°55'N and 116°36’ to 116°50’W, carried out by the O/V Francis- co de Ulloa from CICESE. The material reported was obtained with a Van Veen grab (0.1 m7), during the cruise “BAHIA-10-94” off the Ensenada coast in October 1994. Type specimens are depos- ited in the Polychaetological Collection of the Facultad de Ciencias Bioldgicas, Uni- versidad Autonoma de Nuevo Leon, Méx- ico (UANL); others are in the Centro de Investigacion Cientifica y de Educacién Su- perior de Ensenada, B.C., México (CI- CESE). Nereis fauchaldi, new species Fig. laf Material examined.—Stn. 26 (1) (Holo- type, UANL 3945), [116°44’'N, 31°47'W], 210 m depth. Additional material.—Western coast of Baja California, Sebastian Vizcaino Bay, dredge (6 Jul 1989) Stn. F-10, (1) [28°07'N, 115°00’W], 85 m depth (UANL 3946); Shrimp trawl (6 Jul 1989) Stn. 2 (1) [28°47'N, 114°34'’W], 84 m depth (UANL 3947). Description.—The holotype is an incom- plete specimen, without evident pigmenta- tion pattern, 12 mm long, 1.5 mm wide in- cluding parapodia, with 40 setigers. Prosto- mium pentagonal, two digitate frontal an- tennae, shorter than palps. Eyes large, anterior pair oval, posterior pair round. Palps biarticulate, thin, palpostyles conical. Peristomium longer than next two seg- 824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Nereis fauchaldi, new species: a, anterior end, dorsal view; b, 9th setiger, anterior view; c, 26th setiger, anterior view; d, 39th setiger, anterior view; e, neuropodial heterogomph falciger of setiger 9; f, noto- podial homogomph falciger of setiger 28. Scale bars: a = 1 mm; b—d = 150 pm; e-f = 15 pm. ments, with four short tentacular cirri, lon- cones in 2 oblique rows; III = 14 cones in gest reaching setiger 3 (Fig. la). oval group; IV = 20 cones in crescent Pharynx with brown jaws, each with 6 shape; V = 0; VI = 8 cones in circle; VII— teeth; paragnaths as: I = 2 cones; TH = 12 VIII = 70 small cones in 3 rows near oral VOLUME 111, NUMBER 4 aperture, and 7 larger cones in one distal row. Parapodia of first two setigers uniramus, thereafter biramous. Anterior notopodia with dorsal and median ligule distally con- ical, superior lobe rounded. Neuropodium with postsetal lobe short, end mammili- form, ventral ligule distally rounded. Dorsal cirri inserted medially on dorsal ligule; ven- tral cirri inserted basally (Fig. 1b). Median and posterior parapodia with long dorsal ligules. Median and ventral ligule elongate, narrow, postsetal lobes expanded distally. Dorsal cirri inserted subdistally, ventral cir- ri inserted basally, shorter than dorsal cirri (Fig. lc, d). Anterior notosetae slender homogomph spinigers in supracicular position. Supraci- cular neurosetae similar to notosetae, and a single heterogomph falciger; infracicular setae heterogomph spinigers and falcigers, each falciger with long blade and thin, moderately long serrations, distal part of blade strongly bent (Fig. le). Median and posterior supracicular notosetae homo- gomph falcigers, starting in setiger 14; blade short and distally blunt, without ser- rations; shaft with crenulate distal mem- brane (Fig. 1f). Supracicular neurosetae of median parapodia homogomph spinigers and heterogomph falcigers; infracicular se- tae heterogomph spinigers. Posterior neu- ropodia each with one heterogomph falciger in supracicular position; and one hetero- gomph falciger and spiniger in infracicular position. Pygidium unknown. Remarks.—WNereis fauchaldi, new spe- cies, belongs to the group of species char- acterized by having posterior parapodia with greatly expanded dorsal ligules. This group includes N. angelensis Fauchald, 1972, N. anoculis Hartman, 1960, N. ano- culopsis Fauchald, 1972, N. fossae Fau- chald, 1972, N. heterocirrata Treadwell, 1931, N. ligulata Hilbig, 1992, N. nichollsi Kott, 1951, N. piscesae Blake & Hilbig, 1990, N. profundi Kirkegaard, 1959, N. sandersi Blake, 1985 and N. vexillosa Gru- 825 be, 1851. All of these species, except WN. vexillosa, have been found in deep waters. These species may be separated into two groups: species with dentate homogomph falcigers; and those with smooth homo- gomph falcigers. Nereis fauchaldi, new spe- cies, belongs to the second group, together with N. angelensis and N. fossae. These species differ in paragnath arrangement and insertion of dorsal cirri. The types of Nereis angelensis Fauchald, 1972: holotype (LACM-AHF 1060) and Nereis fossae Fau- chald, 1972: holotype (LACM-AHF 1058), paratypes (LACM-AHF 1059) (12 speci- mens) were reviewed. Nereis angelensis has the following paragnath arrangement: Area I = 2 in line, II = 17 in 3 irregular rows, III = 24 in 4-5 rows, IV = 18-20 in cre- sent shape, V = O, VI = 3 in line right side, 3 in triangle left side, VII-VIII = 35 in one irregular line; NV. fossae has Area I = 2 in line, II = 15 in 2 rows, III = 28 in oval shape, IV = 18 in cresent shape, V = 0, VI = 3 in line, VIJ—-VIII = 6 in one irregular line; N. fauchaldi has Area I = 2 cones, I = 12 cones in 2 rows, III = 14 cones in oval group, IV = 20 cones in cresent shape, V = 0, VI = 8 cones in group, VII-VIII = 70 small cones in 3 rows near the oral ap- erture, and 7 larger cones on one distal row. Nereis angelensis has dorsal cirri inserted basally, while N. fossae has dorsal cirri in- serted medially, and N. fauchaldi has dorsal cirri inserted subdistally. Etymology.—The species is named after Kristian Fauchald in recognition of his valuable contributions to the systematics of the polychaetes of western Mexico. Distribution.—Nereis fauchaldi is known from its type locality, Todos Santos Bay, Baja California, Mexico, and from Sebas- tian Vizcaino Bay, western Baja California. Nereis imajimai, new species Fig. 2a—g Material examined.—Stn. 1, [116°38'N, 31°50’W], 25 m depth, (8 specimens) (UANE 7 73948)) "eStns) 45 te s9iN: 826 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bh oct Se Fig. 2. Nereis imajimai, new species: a, anterior end, dorsal view; b, 10th setiger, anterior view; c, 30th setiger, anterior view; d, 53rd setiger, anterior view; e, notopodial homogomph falciger of setiger 30; f, neuro- podial heterogomph falciger in upper infracicular position of setiger 30; g, neuropodial heterogomph falciger in middle infracicular position of setiger 30. Scale bars: a = 1 mm; b—d = 150 pm; e = 15 pm; fg = 10 pm. 31°47'W], 20 m depth, (3 specimens) m depth, (4 specimens) (CICESE), Stn. 22, (UANL 3951), Stn. 13, [116°40’N, [116°42’N, 31°46’W], 51 m depth, (1 spec- 31°50'W], 24 m depth, (10 specimens) (CI- imen) (CICESE), Stn. 26 (Holotype, VANL CESE), Stn. 14, [117°40’N, 31°51’W], 19 0000), [116°44’N, 31°47’W], 210 m depth. VOLUME 111, NUMBER 4 Additional material.—West coast of Baja California, 4 Sep 1990, stn. H-8, [27°56'N, 114°54’W], 66 m depth (2 specimens) (UANL 3981). Description.—The holotype is an incom- plete specimen, yellowish in color, 15 mm long and 1 mm wide including setae, with 54 setigers. Prostomium longer than wide, with pair of cirriform antennae. Eyes small, anterior pair reniform, posterior pair round. Biarticulated palps, with conical palpostyles. Tentacular ring slightly longer than setiger 1, four pairs of tentacular cirri, longest reaching posteriorly to setiger 5 (Fig. 2a). Pharynx with pair of brown jaws, and with six teeth; paragnaths as: Area I = 0, II = 2 right, 3 left in one row, III = 6 in group, IV = 6 right, 9 left in cresent shape, V = 0, VI = 3 in triangle, VU-—VIII = 55 in 3—4 rows. Setigers 1—2 uniramous, biramous there- after. Anterior notopodia with dorsal and median ligules triangular, subequal; neuro- podia with postsetal lobes distally digiti- form, ventral ligules subulate. Dorsal and ventral cirri inserted basally, similar in length (Fig. 2b). Median and posterior para- podia with dorsal and median ligules thin, triangular in shape, dorsal ones longer. Dor- sal cirri longer than ventral cirri (Fig. 2c, d). Anterior parapodia with supracicular no- tosetae slender homogomph spinigers. Su- pracicular neurosetae heterogomph falci- gers with long finely serrated blades, and homogomph spinigers; infracicular neuro- setae slender heterogomph spinigers and falcigers, falcigers similar to supracicular ones. Middle notopodia with homogomph falcigers, with short blade and four coarse teeth along edge, crenulate membrane along distal margin of shafts (Fig. 2e); supraci- cular neurosetae homogomph spinigers, in- fracicular neurosetae heterogomph falcigers with long blades and fine serrations, similar to those of anterior parapodia (Fig. 2f), and heterogomph falcigers with short blades (Fig. 2g). Posterior parapodia each with no- topodial supracicular homogomph falciger. 827 Supracicular neurosetae homogomph spi- nigers and one heterogomph falciger; infra- cicular setae heterogomph spinigers. Pygidium unknown. Remarks.—Nereis imajimai, new spe- cies, belongs to a small group of species that possesses short dorsal ligules in pos- terior parapodia, homogomph falcigers with short, dentate blades, and no paragnaths on Areas I and V. Other species in this group include N. apalie Wilson, 1985 and N. cir- riseta Hutchings & Turvey, 1982, both known from Australia. N. imajimai differs from these species in paragnath arrange- ment. Nereis apalie has paragnaths only on Areas IV = 4 right, 2 left and Area VI = 1; N. cirriseta has conical paragnaths ar- ranged as: II = 7 in 2 oblique rows, III = 1 minute cone centrally, [V = 9-10 irreg- ularly in oblique crescent shape, VI = 5 in small oval patch, VII-VIII = 7 in single, evenly spaced transverse row. Etymology.—The species is named after Minoru Imajima in recognition of his valu- able contributions on the systematics of the Nereididae. Type locality.—TYodos Santos Bay, Stn. 26 [116°44’'N, 31°47'W], 210 m depth. Distribution.—Nereis imajimai is known from Todos Santos Bay, Ensenada, Baja California, Mexico and the west coast of Baja California. Acknowledgments We wish to express our most sincere gratitude to Dr. Sergio Salazar-Vallejo for his comments and suggestions on an early version of this work. Two anonymous re- viewers made useful comments to improve the manuscript. Literature Cited Berkeley, E., & C. Berkeley. 1958. Some notes on a collection of Polychaeta from northeast Pacific south of Latitude 32°N.—Canadian Journal of Zoology 36:399—407. Blake, J. A. 1985. Polychaeta from the vicinity of deep-sea geothermal vents in the eastern Pacif- ic. I: Euphrosinidae, Phyllodocidae, Hesionidae, 828 Nereididae, Glyceridae, Dorvilleidae, Orbini- idae, and Maldanidae.—Bulletin of Biological Society of Washington 6:67—101. , & B. Hilbig. 1990. Polychaeta from the vi- cinty of deep sea hydrothermal vents in the east- ern Pacific. II. New species and records from the Juan de Fuca and Explorer Ridge system.— Pacific Science 44(3):219—253. Fauchald, K. 1972. Benthic polychaetous annelids from deep water off western Mexico and adja- cent areas in the eastern Pacific Ocean.—Allan Hancock Monographs in Marine Biology 7:1— D5 Grube, A. E. 1851. Annulaten—Jn: Reise in den Au- bersten horden und Osten Sibiriens . Von A. Th. V. Middendorff. Bd. 2, Teil 1, 24 pp., St Pe- tersburg. (not seen) Hartman, O. 1952. Iphitime and Ceratocephala (Poly- chaetous annelids) from California.—Bulletin of the Southern California Academy of Sciences 59:9—20. . 1960. Systematic account of some marine in- vertebrate animals from the deep basins off southern California—Allan Hancock Pacific Expeditions 22:69—216. . 1963. Submarine Canyons of Southern Cali- fornia. 3. Systematics: Polychaetes.—Allan Hancock Pacific Expeditions 27(3):1—93. Hilbig, B. 1992. New polychaetous annelids of the families Nereididae, Hesionidae, and Nephtyi- dae from the Santa Maria Basin, California, with a redescription of Glycera nana Johnson, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1901.—Proceedings of the Biological Society of Washington 105(4):709-722. Hutchings, P A., & A. Reid. 1990. The Nereididae from Australia. Gymnonereididae sensu Fitz- hugh, 1987: Australonereis, Ceratocephale, Dendronereides, Gymnonereis, Nicon, Olganer- eis, and Websterinereis.—Records of the Aus- tralian Museum 42(1):69—100. , & S. P Turvey. 1982. The Nereididae of South Australia.—Transactions of the Royal So- ciety of South Australia 106:93-144. Kirkegaard, J. B. 1959. The Polychaeta of Western Af- rica.—Atlantide Reports 5:7—117. Kott, P. 1951. Nereidae and Eunicidae of South West- ern Australia; also notes on the Ecology of Western Australian limestone reefs.—Journal and Proceedings of the Royal Society of West- ern Australia 35:85—130. Reish, D. J. 1963. A quantitative study of the benthic polychaetous annelids of Bahia de San Quintin, Baja California.—Pacific Naturalist 3:401—436. Treadwell, A. L. 1923. Polychaetous annelids from Lower California with descriptions of new spe- cies.—American Museum Novitatis 74:1—11. . 1931. Three new species of polychaetous an- nelids in the collections of the United States National Museum.—Proceedings of the United States National Museum 80(2):1—5. Wilson, R. S. 1985. Nereis and Platynereis (Polychae- ta: Nereididae) from Victoria with description of a new species of Nereis.—Proceedings of the Royal Society of Victoria 97(3):129-138. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):829—842. 1998. Marphysa belli (Polychaeta: Eunicidae) and two related species, Marphysa oculata and M. totospinata, a new species, with notes on size-dependent features Hua Lu and Kristian Fauchald (HL, KF) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A.; (HL) Department of Biological Sciences, The George Washington University, Washington, D.C. 20052, U.S.A. Abstract.—Three morphologically related species, Marphysa belli, Marphysa oculata and Marphysa totospinata are described or re-described based on a morphometric study. The morphological differences among these species are discussed. The developmental patterns of antennae, branchiae and setae are emphasized. Since the establishment of Marphysa bel- li in 1833 by Audouin and Edwards from the French side of the English Channel, this species has been widely reported from other regions of the world, such as Plymouth, England (McIntosh 1910), New England re- gion of the U.S.A. (Pettibone 1963), and the Gulf of Mexico (Treadwell 1921, Gath- of 1984). Marphysa belli is unique in hav- ing both compound falcigers and compound Spinigers in the anterior region and in hav- ing well-developed branchiae restricted to a short anterior region. With a large number of specimens at hand, this study examines the validity of the cosmopolitan distribution of M. belli and the correlation between the variation of certain morphological charac- ters and body size (width) of the specimens. All specimens were observed with stereo and compound light microscopes. Sketches for the illustrations were made using a cam- era lucida. The statistical figures were pro- duced with the program Origin. For the cor- relation coefficients, the confidence limits were set at 0.05. Marphysa belli Audouin & Edwards, 1833 Figs. la—j, 2—6 Material examined.—Gulf of Mexico, off Florida, USNM 090007 (n = 2). Atlan- tic Coast of U.S.A.: USNM 9166 (n = 1), USNM 28955 (n = 1), USNM 56988 (n = 2), USNM 109810 (n = 2), USNM 109817 (n = 1), USNM 109820 (n = 4), USNM 109823 (n = 1), USNM 109825 (n = 7), USNM 109829 (n = 1), USNM 109830 (n = 2), USNM 109831 (7 = 3), USNM 109838 (n = 1), USNM 109843 (n = 4), USNM 109844 (nv = 3), USNM 120624 (n = 1), USNM 145277 ( = 1), USNM 145280 (n = 1), USNM 145281 (n = 1), USNM 145282 (n = 2), USNM 145283 (n = 1), USNM 145284 @ = 1), USNM 145287 (n = 1), USNM 145290 (n = 1), and USNM 145294 (n = 1). English Chan- nel, France, St. Malo area, Lancieux: NMW.Z.1991.067.2 (n = 2, Natural History Museum of Wales). English Channel, France, Bay of Seine: LAO73A (n = 10, Natural History Museum of Paris) and LAW3A (n = 5, Natural History Museum of Paris). The following description is mainly based on a specimen from the Woods Hole region of Massachusetts (USNM 28955). Abbreviations.—For the prostomial ap- pendages, AI indicates the lateral palps, AII the paired inner lateral antennae and AIII the single median antenna. Species descrip- tions, including the jaw formulae, are pre- 830 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.025mm d.f.a,h,i 0.0imm 0.05mm FESS 9539508 Sag wail ia S953 9 30S5e Das? satin SVAN IES OT WE FY V-tiel 32.00) 6 1 oO DapSsoss~ Fig. 1. Marphysa belli: a, anterior end, lateral view (USNM 28955); b, 14th parapodium; c, 91st parapodium; d, limbate seta; e, pectinate seta; f, compound spiniger; g, compound falciger; h, aciculae; i, subacicular hook; j, maxillae (USNM 109830). VOLUME 111, NUMBER 4 831 14 vO) Vv VV cc) o OvVOYV O wy © Oo te 212 S O68 6 ra) Soe 2 © OKO O) G60 © ies ~ = 10 Go A AA A = ro) dD) A GCOUANOWN AAA A = © A A A re 8 PAN O D © belli — A oculata LL Vv totospinata 0.5 1120) 1.5 2.0 Zoo) 3.0 2.8 Body width, mm (include parapodia) Fig. 2. Size-dependent variation of the first chaetiger with branchiae in M. belli (n = 48), M. oculata (n = 18) and M. totospinata (n = 7). sented in a format similar to that of Fau- chald (1982). Description.—Specimen complete, with 245 chaetigers. Two pairs of anal cirri; dor- sal pair long, reaching last 6 chaetigers. First 10 chaetigers measure 3.0 mm in length, widest region measures 2.3 mm. Prostomium (Fig. 1a) distally rounded and entire, slightly narrower and shorter than, and % depth of, peristomium. Two palps and three antennae, slightly wrinkled, Same size; palps and lateral antennae slight- ly close to each other. Length of AI:AII: All] = 1:1.2:1.5; ATII extending just past anterior end of prostomium; AII reaching edge of anterior end of prostomium. No eyes observed (eyes observed in other spec- imens, see discussion below). First ring of peristomium about % of total peristomial length (dorsal view). No peristomial cirri. First parapodia smaller than others. Dor- sal cirri long throughout body, slender in posterior region. Dorsal cirri always longer than ventral cirri. Postsetal lobe very long in anterior parapodia (Fig. 1b), becoming smaller beginning at first chaetiger (32nd) with subacicular hook (Fig. Ic). Pectinate branchiae present from 12th— 27th parapodia; 12th parapodia with 7 fil- aments, 27th parapodia with 6 filaments, as many as 12 filaments observed on 18th parapodia. Individual branchial filaments longer than dorsal cirri (Fig. 1b). Serrated limbate setae (Fig. 1d) longest of those situated on dorsal side of parapo- dia. Pectinate setae (Fig. le) present from anterior parapodia, with 5—9 teeth; tooth on one edge longer, more stout than others, tooth on other edge slightly longer than oth- ers. One to eight compound spinigers (Fig. 832 50 40 30 20 Last chaetiger with branchiae 10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON © belli A oculata V__ totospinata 0.5 1.0 11.5 2.0 ZS 3.0 3.5 Body width, mm (include parapodia) Fig. 3. Size-dependent variation of the last chaetiger with branchiae in M. belli (n = 49), M. oculata (n = 18) and M. totospinata (n = 7). lf) present from first to about 55th para- podia, situated on posteroventral side of parapodia, with shaft and one side of blade serrated. Compound falcigers (Fig. 1g) present throughout body, arranged in about 3 rows on anteroventral side of parapodia; blade bidentate, with serrated hood and shaft; compound falcigers at least twice as numerous aS compound spinigers in any chaetiger in which both kinds of setae oc- cur. Aciculae (Fig. 1h) 2—3 in number in anterior region, becoming single from an- terior-middle region; honey color, becoming darker towards posterior end; blunt-headed anteriorly, sharper in posterior region. Sub- acicular hook (Fig. 11) bidentate with hood, present from 32nd parapodia, always single, except for replacement; honey to dark hon- ey color. Maxillary formula (Fig. 1j): 1+1, 6+6, 0+6, 8+4, 1+1 (USNM 109820); 1+1, 7+6, 0+6, 8+4, 1+1 (USNM 109830); 1+1, 8+7, 0+8, 9+4, 1+1 (USNM 109831); 1+1, 8+7, O+7, 9+5, 1+) CAWS3A) sal, iano: 0+6, 7+5, 1+1 (LAO73A). Remarks.—The beginning of branchiae (Fig. 2), the ending of branchiae (Fig. 3), the maximum number of branchial fila- ments (Fig. 4), the ending of the compound spinigers (Fig. 5) and the beginning of the subacicular hook (Fig. 6) are all size-de- pendent features (see Table 1). The ending of compound spinigers occurs several chae- tigers posterior to the beginning of the sub- acicular hook. Of 63 specimens examined, only six are complete. Four complete larval specimens, USNM 109818 (49 chaetigers, 0.8 mm VOLUME 111, NUMBER 4 25 NO Oo 833 1) —_ Cc c¢) = acy a oO V © Vv ra O S © 15 Vv Q ©) VV Oo yo © = © G00 © r= 10 (0) © A - oA LAO Cc © AQOAOA = o) © AGE © = C00 GOA A © © belli Sx 5 Loco A Lo A oculata x< NEO) ©) Vv totospinata 40) = 0.5 1.0 159 2.0 29 3.0 S45) Body width, mm (include parapodia) Fig. 4. Size-dependent variation of the maximum number of branchial filaments in M. belli (n = 49), M. oculata (n = 17) and M. totospinata (n = 7). wide), USNM 109817 (70 chaetigers, 0.9 mm wide), USNM 145290 (58 chaetigers, 0.85 mm wide) and USNM 145294 (88 chaetigers, 1.05 mm wide), have only three antennae developed; the lateral palps (AI) have not appeared. Three other incomplete specimens (USNM 56988, 145280, 109820), have a body width of 0.9—1.05 mm and also have only three antennae developed. One complete specimen, USNM 109825 (77 chaetigers, 1.1 mm wide), has three anten- nae and two palps developed; two other small incomplete specimens, USNM 109825 (1.2 mm wide) and USNM 145277 (1.1 mm wide), also have antennae and palps. These observations suggest that M. belli does not develop its palps until it reaches about 80 chaetigers and a body width around 1.1 mm. One pair of very light-colored reniform eyes was observed in most of the specimens from the Atlantic coast of the U.S.A., such as in USNM 109820. For specimens from the French coast of the English Channel, those collected two decades ago, eyes are not observed. All recently collected speci- mens have distinctive eyes. It is likely that the absence of eyes in preserved specimens of M. belli is an artifact of prolonged pres- ervation. When one of us (HL) visited the Natural History Museum in Paris, the type speci- men of M. belli from the Island of Chausey was not available. However, many speci- mens from around the type locality, such as St. Malo, which is in the same bay as the type locality, were examined. Audouin & Edwards’ (1833) original description is co OO TS io) (@) Last chaetiger with compound spinigers PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 60 © © © oO © 50 © 40 Go © oO © O (6) © © belli A oculata 10 ONS (0) 1-5 /ZA0) TZE5 3.0 515) Body width, mm (include parapodia) Fig. 5. Size-dependent variation of the last chaetiger with compound spinigers in M. belli (n = 40) and M. oculata (n = 15). very simple, and it does not contradict the = 2; USNM 130368, n = 6; USNM features we observed in these specimens. Geographic distribution.—M. belli has a wide geographic distribution. It not only occurs on the French side of the English Channel but is also found along much of the Atlantic coast of the U.S.A. (Massachu- setts, Delaware, Georgia) and from one sta- tion in the Gulf of Mexico, off Florida. Marphysa oculata Treadwell, 1921 (Char. emend.) Figs. 2—6, 7a-j, 8 Material examined.—North Atlantic Ocean, Gulf of Mexico, USA, off Florida, Dec 1982 or Jun 1983 (USNM 129779), 25°17'48"N, 81°39'48"W, SOFLA, 14 m, STA 52, n = 12; USNM 129706, n = 2; USNM 130127, n = 1; USNM 130256, n 130484, n = 1. The following description is based mostly on the catalogued specimen USNM 129779. Description.—Specimen complete, with 120 chaetigers. First 10 chaetigers measure 2.1 mm in length, widest region (with para- podia) measures 2.2 mm. Two pairs of anal cirri; long dorsal pair reaching last 5 chae- tigers; small ventral pair only % length of dorsal pair. Prostomium distally round and entire (Fig. 7a); slightly narrower than peristom- ium, subequal in length and less than 4% depth of, peristomium. Three antennae (AII and AIII) and two palps (AD, irregularly slightly wrinkled, evenly spaced, short, similar in thickness. Length of AI:AII:AIII = 1:1.3:1.4. AII and AIII extending to near VOLUME 111, NUMBER 4 45 40 35 30 25 20 15 First chaetiger with subacicular hook 10 OS 140) 18) 835 © belli A oculata Vv totospinata 2.0 P25) 3.0 3.5 Body width, mm (include parapodia) Fig. 6. (n = 17) and M. totospinata (n = 7). anterior end of prostomium. One pair of large, black oval eyes outside of AII and posterior to Al, partly covered by peristom- ium. First ring of peristomium % (dorsally) to % (ventrally) of total peristomial length. No peristomial cirri. First chaetiger equal in length to second ring of peristomium, following chaetigers broader and longer. First parapodia smaller than others. Dorsal cirri long throughout body, always longer than ventral cirri. Post- setal lobe long in anterior parapodia (Fig. 7b), absent from chaetigers with subacicu- lar hook. Ventral cirri digitiform throughout body, becoming smaller from about first chaetiger lacking prominent postsetal lobe. Pectinate branchiae present from 10th to 19th parapodia, with 9 filaments, individual filaments slightly longer than dorsal cirri. Limbate setae (Fig. 7c) longest, finely ser- Size-dependent variation of the first chaetiger with subacicular hook in M. belli (n = 49), M. oculata rated on one edge. Pectinate setae (Fig. 7d) present in anterior body region, with 7-11 teeth, tooth on one edge longer, more stout than others, tooth on other edge slightly lon- ger than others. Compound falcigers biden- tate (Fig. 7e, f), present from 16th parapodia, shaft and base of hood serrated. Compound spinigers (Fig. 7g) serrated on blade and shaft, present on first to 27th parapodia, 20 or more compound spinigers on 10th para- podia. Aciculum (Fig. 7h) with blunt head; yellow to very light honey in color; up to 3 aciculae present from 6th to 8th parapodia, only 1 aciculum present from 14th parapodia to posterior end. Subacicular hook (Fig. 71) bidentate, with hood, distal tooth small and directed obliquely, proximal tooth directed laterally; yellow to very light honey in color; present from 23rd parapodia, always single, except for replacement. 836 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Correlations between body width (X, mm) and the morphometric characters (Y), which is expressed by linear regression ““Y = AX + B,”’ in Marphysa belli (n = 49), M. oculata (n = 17) and M. totospinata (n = 7). SD is standard deviation. R is coefficient. The morphometric characters are: the first chaetiger with branchiae (FB), the last chaetiger with branchiae (LB), the maximum number of branchial filaments (MB), the first chaetiger with subacicular hook (SH), the last chaetiger with compound spiniger (CS) and the first chaetiger with compound falciger (CF). Y A + SD M. belli FB 5.30 + 0.51 M. belli LB 115 2 156 M. belli MB —1.30 + 0.81 M. belli SH 8.46 + 1.22 M. belli CS SESS 243 M. oculata FB TMs = O39) M. oculata LB 10.70 = 0.62 M. oculata MB 1.57 + 0.50 M. oculata SH 13.16 + 0.96 M. oculata CS IES OF=an ES M. oculata CF —5.40 + 1.96 M. totospinata FB 11.90 + 0.63 M. totospinata LB QI 22 Dadi M. totospinata MB 5, 22 JB M. totospinata SH 10.70 + 4.66 Maxillary formula (Fig. 7j): 1+1, 6+5, OF) a4 Lal (U SNM IZ 9779) le 6455, 07-5, 9573.0 1Goleand ae lA 9/-.6505-6; 6+4, 1+1 (USNM 130368). Remarks.—According to morphometric data, the size-dependent morphological var- lation is distinctive in this species. Antennae: we have a complete specimen with 71 chaetigers which possesses three antennae but lacks the lateral pair of palps (AI), whereas several other specimens with about 84 chaetigers possess the lateral palps. These observations suggest that the palps of M. oculata do not develop until it reaches about the 80-chaetiger stage. This estimation is comparable to the palp devel- opmental pattern of Marphysa belli. The palp development of these two species of Marphysa is much later when compared to that of Eunice wui (Lu & Fauchald 1998), whose palps emerge after the worm reaches about 30 chaetigers in size. However, both Eunice and Marphysa of the Eunicidae de- velop their lateral palps much later than species in the Onuphidae. For example, No- tonuphis antarctica (our observations), Nothria elegans (Blake 1975), Mooreonu- B + SD R 3.27 + 0.30 0.85 M23) 32 OG 0.87 5.81 + 0.48 0.87 8.80 + 0.72 0.87 20.60 = 1.43 0.92 1.17 + 0.24 0.78 4.50 + 0.38 0.95 3.46 + 0.31 0.95 4.61 + 0.58 0.90 8.03 = 1.08 0.90 10.96 = 1.21 0.93 0.63 + 0.23 0.78 S29 50s 2103 0.89 3.51 + 0.48 0.96 QQ) 22 il 0.92 phis jonesi (Fauchald 1982) and Kinber- gonuphis simoni (Hsieh & Simon 1987) have developed the lateral palps when they are only 5 chaetigers in size. It appears that the pattern of development of the palps is more uniform among the Onuphidae when compared to the Eunicidae. Branchiae: before the development of the palps, the branchiae are already well devel- oped. During the development of the worm, the starting chaetiger with branchiae chang- es from the 8th parapodia to the 10th para- podia (Fig. 2, Table 1); the ending chaetiger with branchiae changes from the 13th para- podia to the 21st parapodia (Fig. 3, Table 1); the number of branchiae changes from only 6 pairs to 12 pairs; the maximum num- ber of branchial filaments changes from 4 to 10 (Fig. 4, Table 1). Compound falcigers (Fig. 8, Table 1): their number is always less than that of the compound spinigers in any given segment in which both occur. In young worms (0.7 mm wide), compound falcigers are present in all chaetigers. When the worm grows, compound falcigers are lost from a certain number of anterior chaetigers. The loss VOLUME 111, NUMBER 4 837 imm a BNNs | uN \ WS \ 3 = 0.1mm i Resse b _~ 0.0 m 0.01MM « ge,f.g,hi SID Fig. 7. Marphysa oculata: a, anterior end, lateral view (USNM 129779); b, 15th parapodium; c, limbate seta; d, pectinate seta; e, f, compound falcigers; g, compound spiniger; h, aciculum; i, subacicular hook; j, maxillae. 83 oo ss — Ke) (e) on Oo First chaetiger with compound falciger 0.5 1140) Heo PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2.0 25 Body width, mm (include parapodia) Fig. 8. Starts at about chaetiger 8 and proceeds from there in both directions. For a 2.5 mm wide worm, compound falcigers are absent in the first 19 parapodia. Compound spinigers (Fig. 5, Table 1): the ending position of the compound spi- nigers is size-dependent and varies from the 16th chaetiger to the 31st chaetiger. Subacicular hook (Fig. 6, Table 1): the first occurrence of a subacicular hook changes from the 15th parapodia to the 24th parapodia. It always occurs several chaetig- ers after the ending of the branchiae and several chaetigers before the ending of the compound spinigers. The sensory organ reported by Hayashi & Yamane (1994) for Marphysa sanguinea is also observed on the ventral base of dor- sal cirri (22rd parapodia) of M. oculata (USNM 130127). Size-dependent variation of the first chaetiger with compound falcigers in M. oculata (n = 15). Treadwell (1921) described a single in- complete specimen from Key West Harbor, Florida (close to our collection site) as Mar- physa belli, variety oculata based on the presence of eyes in his specimen versus the absence of eyes in European specimens, though he mentioned with contradiction that ‘‘Since the presence or absence of eyes does not in itself seem to me of specific importance, I have regarded this as merely a varietal difference.’ As indicated earlier, the eye condition of M. belli is likely relat- ed to preservation. Treadwell’s description on M. belli, variety oculata totally agrees with our specimens, especially for the setal distribution, i.e., compound spinigers are the main compound setae in the anterior re- gion. Thus, we believe that they belong to the same species. Unfortunately, Treadwell (1921) did not mention the consistent dif- VOLUME 111, NUMBER 4 ferences between his species and M. belli. In the anterior body region, there are at least twice as many compound falcigers than compound spinigers in M. belli, whereas in M. oculata the compound spi- nigers are at least twice as common as the compound falcigers, or the compound fal- cigers may be absent in some anterior para- podia. Only after the observation of many specimens (M. oculata, n = 24; M. belli, n = 63) did this morphological difference be- tween the two species become obvious, thereby warranting a separate species status for M. oculata. Marphysa totospinata, new species Figs. 2—4, 6, 9a—h Material examined.—Atlantic coast, Ire- land, Mayo, near Blacksod Bay (Holotype: NMW.Z.1988.069.84, and 6 paratypes: NMW.Z.1988.069.84 and NMW.Z.1988. 069.84). Description.—Specimen complete, with 322 chaetigers. Two pairs of anal cirri; lon- ger dorsal pair reaching last 4 chaetigers. First 10 chaetigers measure 3.4 mm in length, widest region measures 5.9 mm. Prostomium distally round and entire (Fig. 9a), slightly narrower than, same length and % depth of, peristomium. Three antennae (AII and AIII) and two palps (AD, slightly irregularly wrinkled, same size, evenly spaced; length of AI:AII:AIII = 1: 1.2:1.4; all extend past anterior end of pro- stomium, AI reaching first ring of peristom- ium, AII reaching second ring of peristom- ium, AIII reaching anterior end of first chaetiger. One pair of light brown reniform eyes between AI and AII. First ring of peri- stomium about % of total length (dorsal view). No peristomial cirri. Anterior four chaetigers thick and cube- Shaped; becoming flatter dorsoventrally from fifth chaetiger. First parapodia smaller than others. Dorsal cirri long, finger-shaped from beginning, becoming slender in pos- terior region; dorsal cirri always longer than ventral cirri. Ventral cirri with oval base in 839 anterior chaetigers, becoming slender pos- teriorly. Postsetal lobe long in anterior para- podia until about 65th parapodia, length comparable to dorsal and ventral cirri in some chaetigers. Sensory organ similar to that described by Hayashi & Yamane (1994) for Marphysa sanguinea and M. oculata also observed on ventral base of dorsal cirri of 286th parapodia (Fig. 9b). Pectinate branchiae present from 14th— 40th parapodia. Twelveth parapodia with 11 filaments, as many as 18 filaments observed on 27th parapodia. Individual branchial fil- aments slightly longer than dorsal cirri. Branchiae completely cover dorsal region of worm. Serrated limbate setae longest of those situated on dorsal side of parapodia. Pecti- nate setae (Fig. 9c) present from anterior parapodia; with 8—9 teeth, tooth on one edge longer, more stout than others, tooth on other edge slightly longer than others. Compound falcigers (Fig. 9d) present throughout body, with serrated hood and serrated shaft; more than 20 compound fal- cigers arranged in several rows on antero- ventral side of parapodia, fewer in number on posterior side. Compound spinigers (Fig. 9e) present from beginning till very poste- rior end, such as on parapodium 286 (Fig. 9b); serrated on shaft and one side of blade; 6—7 compound spinigers arranged in a line on posteroventral side of parapodia, de- creasing to 1—2 in number in posterior re- gion. Compound falcigers at least twice as numerous as compound spinigers in ante- rior parapodia. Aciculae (Fig. 9f) 3—4 in an- terior region, single or occasionally double from middle-anterior region; honey color, becoming darker in posterior region; blunt- headed, with sheath. Subacicular hook (Fig. 9g) from 46th parapodia, single in 75% of parapodia, double in those remaining; hon- ey to dark honey color; bidentate with hood. Maxillary formula (Fig. 9h): 1+1, 7+6, 0+7, 8+5, 1+1 in one paratype (NMW.Z. 1988.069.86). Remarks.—The beginning (Fig. 2) and 840 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ~ = Sensory Organ 0.1mm 0.5mm b, h —e———— 4 C, d, e, it g arse c d e f g Fig. 9. Marphysa totospinata, new species: a, anterior end, dorsal view (NMW.Z.1988.069.84); b, 286th parapodium; c, pectinate seta; d, compound falciger; e, compound spiniger; f, aciculum; g, subacicular hook; h, maxillae (NMW.Z.1988.069.86). VOLUME 111, NUMBER 4 ending (Fig. 3) position of branchiae, the maximum number of branchial filaments (Fig. 4) and the starting position of the sub- acicular hook (Fig. 6) of M. totospinata are all size-dependent characters (see Table 1). This species (7 = 7) is consistently differ- ent from M. belli (n = 63) and M. oculata (n = 24) in its setal arrangement: in the anterior parapodia, compound spinigers are the dominant setal type in M. oculata, whereas in M. totospinata compound fal- cigers are more numerous; in M. belli, com- pound spinigers occur on less than %4 of the anterior parapodia, whereas in M. totospi- nata compound spinigers are present nearly throughout the entire body. McIntosh (1910) described a specimen of M. belli from the Plymouth region. Accord- ing to his description, compound spinigers exist from the beginning to the posterior end of the worm. If this is the case, his specimen should belong to M. totospinata. Etymology.—The species is named for its special morphological character, which is the occurrence of compound spinigers throughout the body length. Discussion The branchial and setal patterns are size- dependent morphological characters in Marphysa belli and its two related species, M. oculata and M. totospinata. Among the 15 sets of correlations calculated at a con- fidence limit of 0.05 (Table 1), the range of coefficients (R) is between 0.78 and 0.96 (average = 0.89). The general morpholog- ical variations fall in a similar range be- tween M. belli and M. oculata and between M. belli and M. totospinata, for a similarly sized specimen. Slight morphometric dif- ferences exist between M. oculata and M. totospinata. For example, in a specimen of M. oculata that is 2.5 mm wide, its bran- chiae begin on the 10th parapodia and end on the 22nd parapodia, and its subacicular hook emerges from the 23rd parapodia, whereas for a similarly sized specimen of M. totospinata, its branchiae begin on the 841 13th parapodia and end on the 32nd para- podia, and its subacicular hook emerges from the 32nd parapodia. Though only rel- atively small sized specimens of M. oculata and large sized specimens of M. totospinata were available for the present study, several specimens of both species have overlapping size ranges. In addition, both species show distinctive features when compared to sim- ularly sized specimens of M. belli, whose size ranged from small (larva) to large specimens. These observations support the suggestion that M. oculata is not the juve- nile stage of M. totospinata. The arrange- ment of compound setae is the major dif- ference among these species. If additional specimens of M. belli from other areas of the world, such as the Mediterranean, South Africa and Indochina, can be examined, it is possible that new sister species can be separated. This species group potentially provides a good case for further biogeo- graphic study. Key to the Marphysa belli group 1. At least twice as many compound fal- cigers than compound spinigers in an- tenon paranodal aaa ome a nae 2 1’. At least twice as many compound spi- nigers than compound falcigers in an- teHORpakapoGiayer wee M. oculata 2. Compound spinigers limited to anterior ¥% body region or less .......... M. belli 2'. Compound spinigers present along near- lyathe entire body 4 4-45- M. totospinata Acknowledgments The authors would like to thank Jon No- renburg, Frederick Pleijel, Linda Ward, Cheryl Bright, Bill Moser and other col- leagues from the Division of Worms, NMNH, Smithsonian Institution, and Diana Lipscomb and the Systematic Group, De- partment of Biological Sciences, George Washington University. We thank Dr. An- drew Mackie (Wales) for lending us speci- mens and Dr. Jean-Claude Dauvin (France) for arranging a visit for HL. Stephen Gar- diner and an anonymous reviewer helped to 842 improve the quality of this paper. HL is supported by the Research Enhancement Funds and Weintraub Research Fellowship from the George Washington University. Literature Cited Audouin, J. V., & A. Milne Edwards. 1833. Recherch- es pour servir a lhistoire du littoral de la France. Annelides.—Paris t.2:1—290. Blake, J. A. 1975. The larval development of poly- chaeta from the northern California coast. II. Nothria elegans (Family Onuphidae).—Ophelia 13:43-61. Fauchald, K. 1982. Description of Mooreonuphis jo- nesi, a new species of onuphid polychaete from shallow water in Bermuda, with comments on variability and population ecology.—Proceed- ings of the Biological Society of Washington 95:807—-825. Gathof, J. M. 1984. Eunicidae. Pp. 40.1—40.31 in J. M. Uebelacker & P. G. Johnson, eds., Taxonom- ic guide to the polychaetes of the northern Gulf PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of Mexico, vol. 6, Chapter 40. Barry A. Vittor and Associates, Inc. Mobile. Hayashi, I., & S. Yamane. 1994. On a probable sense organ newly found in some eunicid poly- chaetes.—Journal of the Marine Biological As- sociation 74:65—770. Hsieh, H. L., & J. Simon. 1987. Larval development of Kinbergonuphis simoni, with a summary of development patterns in the family Onuphidae (Polychaeta).—Proceedings of the Biological Society of Washington 7:194—210. Lu, H., & K. Fauchald. 1998. Description of Eunice weintraubi and E. wui, two new species of eu- nicid polychaetes from northern Gulf of Mexi- co.—Proceedings of the Biological Society of Washington 111:230—240. McIntosh, W. C. 1910. The British Annelids. Vol. II, pt. II, Polychaeta, Syllidae to Ariciidae. Pp. 233-524. Ray Society Monographs. Pettibone, M. H. 1963. Marine polychaete worms of the New England region. 1. Aphroditidae through Trochochaetidae.—Bulletin of the Unit- ed States National Museum 227(1):1—356. Treadwell, A. L. 1921. Leodicidae of the West Indian region.—Carnegie Institution of Washington (XV):1-131. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):843-848. 1998. Pseudechiniscus asper, a new Tardigrada (Heterotardigrada: Echiniscidae) from Hokkaido, northern Japan Wataru Abe, Kazuo Utsugi, and Masatsune Takeda (WA) Department of Biological Sciences, Graduate School of Science, The University of Tokyo. (Corresponding address: Department of Zoology, National Science Museum, 3-23-1 Hyakunincho, Shinjuku-ku, Tokyo, 169-0073 Japan); (KU) Hiraoka Environmental Science Laboratory, 8-8 Hosoyama, Asao-ku, Kawasaki-shi, Kanagawa, 215-0001 Japan; (MT) Department of Zoology, National Science Museum, 3-23-1 Hyakunincho, Shinjuku-ku, Tokyo, 169-0073 Japan Abstract.—A new semi-terrestrial tardigrade species of the family Echinis- cidae, Pseudechiniscus asper, is described from a single male collected from Hokkaido, northern Japan. This species can be readily distinguished from the known congeners by a combination of the following characters: almost the entire surface of all dorsal plates is covered with coarse cuticular granulation; the subtriangular projection is present at position D; paired lobes on pseudoseg- mental plate are roundly triangular and tipped with papillate projection; and, each internal claw is armed with a spur. In autumn of 1988, the second author collected lichen samples to obtain the tar- digrades around Lake Shikotsu in south- western Hokkaido, northern Japan. Tardi- grade specimens extracted from the lichen samples were fixed with 5% formalin, and mounted on glass slides with Hoyer’s me- dium. Cover slips were sealed by Paraffin- Balsam. Close examination was performed using a phase contrast and Nomarski dif- ferential interference microscope (ZEISS Axiophoto) and illustrations were depicted with the aid of camera lucida. After a review of the literature, it was concluded that one of the specimens rep- resents a new species belonging to the ge- nus Pseudechiniscus of the family Echin- iscidae, which is described and illustrated as Pseudechiniscus asper, new species. Ter- minology is mainly that used in Dastych & Kristensen (1995) and Kristensen (1987). Pseudechiniscus asper, new species Figs. 1-2 Type material.—Holotype: Four-clawed adult male; Shikotsuko-onsen, Chitose, southwestern Hokkaido, northern Japan (141°24’N, 42°46’E); ca. 250 m; extracted from a foliose lichen, Phaeophyscia imbri- cata, on trunk of Acer japonicum, 9 Oct 1988, K. Utsugi leg. Holotype is deposited in the National Science Museum, Tokyo (NSMT), under the registered number NSMT-Tg 44. Diagnosis.—Small size. Dorsal plates covered with coarse cuticular granulation. Scapular plate and segmental paired plates II and III each with segmental lateral plates. Subtriangular projection at position D. Paired lobes on pseudosegmental plate roundly triangular, tipped with papillate projection. Internal claws each with a spur. Description.—Body length 166 wm ex- cluding leg IV, width 83 wm; body width: body length, 1:2. Eyespot not detected, body color evenly translucent in Hoyer’s medium. Dorsal plates well developed and con- spicuous. Delineation of dorsal plates and subdivision within a plate each comprised of cuticular ridge. Almost entire surface of 844 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 50um 10um cD Fig. 1. cS D Pseudechiniscus asper, new species, holotype male (NSMT-Tg 44), Hokkaido, N Japan: A, Habitus, dorsal view; B, Habitus, ventral view; C, Internal claw of leg II; D, External claw of leg Il. Abbreviations: cr = cuticular ridge; D = subtriangular projection at position D; HP = head plate; ILPI, ILPII = intersegmental lateral plates I-II; MPI, MPII, MPIII = median plates I-III (a: anterior, b: posterior); NP = neck plate; PPI, PPIII = segmental paired plates II-III; PSP = pseudosegmental plate; ScLP = scapular segmental lateral plates; SeLPII, SeLPII = segmental lateral plates II-III; SP = scapular plate (a: anterior, b: posterior); TP = terminal plate. all dorsal plates covered with coarse, uni- formly distributed granules which consist of hemispherical cuticular thickenings; poste- rior half of head plate and neck plate only with very fine, densely distributed puncta- tions which consist of pillar structures of epicuticle; and small region around primary papilla and cirrophore of cirrus A devoid of any sculpture. Granules larger on segmental paired plates II and III, and pseudosegmen- tal plate, ca. 1.5 1m in diameter; granules medium in size on anterior half of head VOLUME 111, NUMBER 4 845 Fig. 2. Pseudechiniscus asper, new species, holotype male (NSMT-Tg 44), Hokkaido, N Japan, differential interference contrast photomicrographs: A, Habitus, dorsal view; B, Internal claw of leg II; C, Detail of segmental paired plate III. Scales = 50 pm (A), 10 pm (B, C). plate, scapular plate, median plates, and ter- minal plate, ca. 0.9 4m in diameter; gran- ules smaller on most anterior part of scap- ular plate, and intersegmental lateral plates I and II, ca. 0.5 pm in diameter. Granules larger in central region of each plate than those in peripheral region of same plate. Space between granules smooth, without striae. Head plate (HP) well-defined, gently fac- eted, subdivided into smaller pieces by smaller anterior and larger posterior W- shaped sculptures; ridge of large W-shaped sculpture effaced in its middle. Most pos- terior points of large W-shaped sculpture and minuscule indentations of neck plate interconnected by rather shallow, incon- spicuous cuticular furrow which is dilated anteriorly. Neck plate (NP) narrow but appreciable; almost entire neck plate overlapping most posterior part of head plate; posterior edge of neck plate with a pair of vague, minus- cule indentations. Scapular plate (=plate I; SP) well devel- oped, inconspicuously subdivided at mid- dorsal line of body, shallowly subdivided into anterior and posterior subrectangular parts (SPa, SPb); anterior part fairly larger than posterior part. Three pairs of scapular segmental lateral plates (ScLP) separated from central main part; anterior segmental lateral plate connected with anterior central part; two pairs of posterior segmental lateral plates arranged transversely, connected with posterior central part. Median plate I (MPI) distinctly subdivid- ed into anterior and posterior parts (MPIa, 846 MPIb); anterior part subrectangular, fairly larger, somewhat wider than posterior part, inconspicuously but completely subdivided at middorsal line of body; posterior part separated from anterior part as a transverse strip, concave dorsally, undivided at mid- dorsal line of body; anterior and posterior edges of median plate I slightly and mod- erately angulated at middorsal line of body respectively. Intersegmental lateral plate I (ILPI) sub- divided into anterior and posterior parts; an- terior part wider than posterior part; proxi- mal edge of two parts obtuse. Segmental paired plate II (PPID) well de- veloped, with two pairs of lateral segmental plates (SeLPII); posterior segmental lateral plate somewhat larger than anterior seg- mental lateral plate; central main part of segmental paired plate II with cuticular ridges (cr) laterally; ridges effaced centrally forming separate anterior and posterior lines. Cuticular ridge, which is made up by posterior edge of segmental plate II and an- terior edge of intersegmental lateral plate II, forming blunt swelling in its most distal part. Median plate II (MPII) larger than me- dian plates I and III, distinctly subdivided into anterior and posterior parts (MPIla, MPIIb); anterior part fairly larger and wider than posterior part, weakly subdivided at middorsal line of body; posterior part form- ing a narrow transverse strip, concave dor- sally, undivided at middorsal line of body; anterior and posterior edges of median plate II angulated at middorsal line of body. Intersegmental lateral plate If (LPI) sub- divided into anterior and posterior parts; an- terior part larger than posterior part; prox- imal points of two parts acute. Segmental paired plate III (PPIII) well developed, similar to segmental paired plate II in size and shape, with two pairs of lat- eral segmental plates (SeLPIII); anterior and posterior lateral segmental plates in- completely separated; central main part of segmental paired plate III with cuticular ridges (cr) laterally; ridges effaced centrally PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON forming separate anterior and posterior lines. Projection at position D (D) subtrian- gular, broad-based, ca. 9 wm long. Median plate III (MPIII) well-defined, rhomboid, undivided; anterior and posterior edges of median plate III moderately and poorly angulated at middorsal line of body, respectively. Pseudosegmental plate (PsP) distinctly and wholly subdivided at middorsal line of body; paired lobes roundly triangular, each tipped with papillate projection, 8 wm high, 17 wm wide; lobes slightly conjoined mid- dorsally with each other; posterior edge of pseudosegmental plate weakly arcuate but not sinuate. Posterior edge of pseudoseg- mental plate covering anterolateral side of terminal plate swollen in appearance. Terminal plate (TP) with remarkable, long cuticular ridge posterolaterally, thus having a strongly facetted appearance, and with very blunt cuticular ridge on its mid- dorsal line of body. Subcephalic plate thin but comparably conspicuous, situated below mouth open- ing; anterior edge moderately marked but effaced in its middle, distinctly arcuate; lat- eral edge well marked, arcuate, dilated an- teriorly; posterior edge scarcely marked, ar- cuate. Dense patches of cuticular granula- tion situated between each pair of legs I- III, and circum-genital region; each granule similar to that on dorsal plates, but much smaller; granules on central part of each patch somewhat larger than those on pe- ripheral part; patch above gonopore with weakly developed edge, thus having a plate-like appearance, other patches without perceptible edge; patches between legs in- creasing in size posteriorly. Net pattern on ventral surface below subcephalic plate throughout but barely visible. Leg plates developed on outer surfaces of legs I-III and dorsal surface of leg IV; each leg plate covered with granules similar to those on dorsal plates; granules larger but sparse on central part of leg plate, smaller but dense on peripheral part. Plate of leg IV VOLUME 111, NUMBER 4 without dentate collar. Sensory organ ab- sent from legs I-III, not detected on leg IV. Internal and external cirri and secondary clava situated ventrally. Internal and exter- nal cirri filamentous, without true cirro- phore but with swollen base, without an- chor-shaped bifurcation at tip; external cir- rus longer than internal cirrus, 12 wm long and 8 pm long, respectively. Secondary cla- va hemispherical, situated nearer to external cirrus than to internal cirrus. Primary clava and cirrus A situated an- terolaterally on scapular plate; primary cla- va papillate, situated just beneath base of cirrus A, ca. 1.5 wm long; cirrus A fila- mentous, short, with normal subconical cir- rophore but without bifurcation at tip, ca. 20 wm long, 12% of body length. Claw normally curved distally, thickened basally; internal claw slightly longer than external claw on all legs; claws of leg I-III ca. 9 wm long; claws of leg IV longer than claws of other legs, ca. 11 pm. External claw on all legs without spur; each internal claw on all legs with a distinct, moderately curved spur directed toward base of claw; spur located at a height approximately % as long as claw length; apex of spur not reach- ing base of claw. Mouth opening situated ventrally. Buccal tube and stylet support weakly sclerotized, thus hardly discernible. Pharyngeal bulb ovoid, 14 pm long, 11 wm wide; outer cu- ticular lining inside pharyngeal bulb clearly thicker than bar-shaped placoid. Male gonopore simple, being a slightly raised ring; anus normal, situated between leg IV of both sides. Etymology.—From the Latin word asper (rough), referring to the general appearance of dorsal plates in the present new species. Remarks.—In Pseudechiniscus asper, de- scribed in this paper, the single specimen collected is male. Pseudechiniscus males are very rare and have been reported only two times, viz. Dastych (1987) as P. alberti and Kristensen (1987) as unidentified spe- cies. Kristensen (1987) stated that Pseude- chiniscus species with males are found 847 limnic or in other stable biotopes and that this is related to the thin sclerotization of the dorsal plates and the absence of the leg plates. It might be true in the species stud- ied by Kristensen (1987), but it is remark- able that both P. alberti from Canada and P. asper from Japan were found in unstable environments such as terrestrial moss or li- chen and have thick dorsal plates. Addi- tional data are required to reveal the rela- tionship between environmental habitat and the mode of reproduction. Kristensen (1987) stated that all Pseu- dechiniscus species are provided with the sensory organ on leg I. However, there are some species lacking this character, e.g., P. Juanitae in Maucci (1986), P. suillus in Ra- mazzotti & Maucci (1983), Dastych (1989), and Kathman & Dastych (1990), P. victor in Maucci (1986), and P. cfr. suillus in McInnes (1995). Also, in P. asper the sen- sory organ on leg I is absent. The sensory organ on leg IV, which is usually developed as a papilla or spine on its outer surface in Pseudechiniscus (Kris- tensen 1987), is not detected in the new species. This character is sometimes hard to observe, depending on the direction of the specimen on the glass slide, and we could not decide whether this character is absent or present in P. asper. The large size of granules on dorsal plates, the presence of a subtriangular pro- jection at the position D, and the presence of segmental lateral plates in the scapular plate and segmental paired plates II-III dis- tinctly separate the new species from spe- cies such as Pseudechiniscus brevimontan- us Kendall-Fite & Nelson, 1996, P. nataliae Biserov & Maucci in Biserov, 1986, P. ra- mazzottii Maucci, 1952, P. scorteccii Fran- ceschi, 1952. P. asper bears some resem- blance to P. occultus Dastych, 1980 from Poland, especially in the general appear- ance of the granulation on the dorsal plates, but can be readily distinguished from it mainly by the absence of the posterior lobe on the scapular plate, the absence of a spurs on the external claws of leg IV, and the ab- 848 sence of anchor-shaped bifurcations of ce- phalic cirri and cirrus A. The last character was omitted in the original description of P. occultus (Dastych 1980) but later emend- ed as presented in Dastych (1987, 1988). Acknowledgments We would like to express our gratitude to Dr. Hieronymus Dastych of the Zoologisch- es Institut und Zoologisches Museum der Universitat Hamburg for critically reading the first draft of this manuscript, and to Mr. Yoshihito Ohmura of the Graduate School of Science, The University of Tokyo for identifying the lichen sample. We are also grateful to anonymous reviewers who kind- ly made invaluable suggestions. Literature Cited Biserov, V. I. 1986. Terrestrial water bears from the North Caucasus. 1. Heterotardigrada.—Zoolo- gicheskii Zhurnal 65:747—756. [In Russian with English summary]. Dastych, H. 1980. Niesporczaki (Tardigrada) Tatrzan- skiego Parku Narodowego. [Tardigrades from the Tatra National Park].—Monografie Fauny Polski 9:1—232. [In Polish with English sum- mary]. . 1987. Two new species of Tardigrada from the Canadian Subarctic with some notes on sex- ual dimorphism in the family Echiniscidae.— Entomologische Mitteilungen aus dem Zoolo- gischen Staatsinstitut und Zoologischen Muse- um Hamburg 8:319—334. . 1988. The Tardigrada of Poland.—Monografie Fauny Polski 16:1—255 + pls. 1-31. . 1989. An annotated list of Tardigrada from the Antarctic—Entomologische Mitteilungen PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON aus dem Zoologischen Museum Hamburg 9: 249-258. , & R. M. Kristensen 1995. Echiniscus ehren- bergi sp. n., a new water bear from the Hima- layas (Tardigrada).—Entomologische Mitteilun- gen aus dem Zoologischen Museum Hamburg 11:221—230. Franceschi, T. 1952. Sul ritrovamento in Valcamonica di Pseudechiniscus novaezeelandiae f. marinae Bartos e di Pseudechiniscus scorteccii n. sp. (Tardigrada).—Doriana 1(30):1—7. Kathman, R. D., & H. Dastych. 1990. Some Echinis- cidae (Tardigrada: Heterotardigrada) from Van- couver Island, British Columbia, Canada.—Ca- nadian Journal of Zoology 68:699—706. Kendall-Fite, K., & D. R. Nelson 1996. Two new spe- cies of tardigrades from Short Mountain, Ten- nessee, USA.—Zoological Journal of the Lin- nean Society 116:205—214. Kristensen, R. M. 1987. Generic revision of the Echin- iscidae (Heterotardigrada), with a discussion of the origin of the family. Pp. 261—335 in R. Ber- tolani, ed., Biology of Tardigrades: Proceedings of the 4th International Symposium on the Tar- digrada, Modena, September 3-5, 1985.—Col- lana U.Z.I. Selected Symposia and Monographs 1, Mucchi Editore, Modena, 380 pp. Maucci, W. 1952. Un nuovo Pseudechiniscus del Car- so Triestino (Tardigrada, Scutechiniscidae).— Atti della Societa Italiana di Scienze Naturali e del Museo Civico di Storia Naturale in Milano 91:127—130. . 1986. Tardigrada.—Fauna d’Italia 24, Edi- zioni Calderini, Bologna, 388 pp. McInnes, S. J. 1995. Tardigrades from Signy Island, South Orkney Islands, with particular reference to freshwater species; Signy Island; South Ork- ney; Antarctica—Journal of Natural History 29:1419-1445. Ramazzotti, G., & W. Maucci 1983. Il philum Tardi- grada: III edizione riveduta e aggiornata.—Me- morie dell’ Istituto Italiano di Idrobiologia 41:1— 1012. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):849—856. 1998. Parapetalophthalmus suluensis, a new genus and species (Crustacea: Mysidacea: Petalophthalmidae) from the Sulu Sea Masaaki Murano and Manuel Rafael Bravo (MM) Institute of Environmental Ecology, METOCEAN Co. Ltd., Riemon 1334-5, Ooigawa-cho, Shida-gun, Shizuoka 421-0212, Japan; (MRB) Vazquez Lopez, 45, 2°, Izquierda, 21001 Huelva, Spain Abstract.—A new genus, Parapetalophthalmus, is established for the new species P. suluensis, collected from the Sulu Sea. It is closely related to Pseu- dopetalophthalmus, but easily distinguished from this genus by the sixth to eighth thoracopods in which the endopods are small, 1- or 2-segmented rudi- ments. During a cruise to southeastern Asian seas (KH-72-1) by the R/V Hakuho Maru of the Ocean Research Institute, University of Tokyo, mysid specimens belonging to the family Petalophthalmidae were collect- ed from the Sulu Sea. They resemble spe- cies of the genus Pseudopetalophthalmus in appearance, but are distinctly different from it by the rudimentary endopods on the sixth to eighth thoracopods. Parapetalophthal- mus Suluensis, new genus, new species, is established for these specimens. The type specimens are deposited in the National Science Museum, Tokyo (NSMT), Japan. Parapetalophthalmus, new genus Diagnosis.—Body slender. Carapace very short, leaving posterior 5 or 6 thoracic somites exposed dorsally; rostrum very short, rounded, without acute process, flanked by pair of small denticles. Eye de- veloped and elongate; cornea spherical, broader than stalk; eyestalk elongate, with- out papilla. Antennular peduncle very long, slender. Antennal scale lanceolate, proximal *2 of outer margin naked, ending in 1 joint- ed spine, distal % of outer margin setose; apex and inner margin setose. Mandibular palp long, powerful and prehensile; third segment short and robust, with 9 long spines. Thoracopods: first pair without ex- opod, endopod terminating in claw, with in- ner lobe on ischium; second pair without exopod, endopod robust, terminating in strong, curved claw, with quadrangular, large, inner lobe on ischium; third to fifth pairs with endopod slender, without termi- nal claw; sixth to eighth pairs with endopod rudimentary. Pleopods of male biramous, natatory; of female biramous, exopod very slender, endopod fused with sympod. Uro- pod: exopod 2-segmented, proximal seg- ment with outer margin naked except for 2 spines and | seta at distal end, margin of distal segment setose; endopod without spines but with setae, statocyst absent. Tel- son rectangular; apex with 9 spine groups and 8 plumose setae between them, each spine group composed of 3 spines with middle spine longest, spines of outermost group barbed, longer, stouter, articulated at base. Type species.—Parapetalophthalmus su- luensis, new genus, new species. Etymology.—The generic name is de- rived from the close resemblance to Pseu- dopetalophthalmus and Petalophthalmus; it is a masculine gender. Remarks.—The new genus is closely re- 850 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Morphological differences between Pseudopetalophthalmus Bravo & Murano, 1997, and Parape- talophthalmus, new genus. Pseudopetalophthalmus Eyestalk Third segment of mandibular palp Endopods of 3rd to 5th thoracopods Endopods of 6th to 8th thoracopods lated and morphologically similar to Pseu- dopetalophthalmus, which was recently es- tablished by Bravo & Murano (1997). The most distinctive difference between the two genera is found in the thoracopods. In Pseu- dopetalophthalmus the endopods of the third to eighth thoracopods are long, multi- segmented, and bearing a terminal claw, whereas in the new genus those of the sixth to eighth thoracopods are small, 1- or 2- segmented rudiments, and those of the third to fifth thoracopods terminate in 2 setae but not in a claw. Parapetalophthalmus and Pseudopetal- ophthalmus can also be separated by dif- ferences in the eyestalk and mandibular palp. In the new genus the eyestalk is not armed with a papilla, and the third segment of the mandibular palp is armed with 9 long spines as compared to 7 in Pseudopetaloph- thalmus. Morphological differences be- tween Pseudopetalophthalmus and Para- petalophthalmus are summarized in Table 1. The new genus also resembles Petaloph- thalmus, but is easily distinguished from it by characteristics of the rostrum, antennal scale, mandibular palp, thoracic endopods, and telson. Parapetalophthalmus suluensis, new species Figs. 1-4 Type specimens.—Holotype (NSMT-Cr 12168), adult male (13.0 mm); allotype (NSMT-Cr 12169), adult female (11.4 mm); paratypes (NSMT-Cr 12170), 1 immature female (9.1 mm), 1 juvenile (5.2 mm); Sulu Sea, 08°12.7'N, 117°59.6’E to 08°11.8'N, With small papilla With 7 long and strong setae Terminating in a strong claw Long and multi-segmented Parapetalophthalmus Without papilla With 9 long and strong setae Terminating in 2 short setae Reduced to 1- or 2-segmented, small rudiment 117°58.4'E, 285-306 m, plankton net in- stalled in mouth of 3-m beam trawl, 26 May WZ: Description.—Body slender. Carapace very short, leaving posterior 5 or 6 thoracic somites exposed dorsally; frontal margin broadly rounded, upturned, not forming def- inite rostral projection, leaving eyes com- pletely uncovered; small blunt process on either side of margin (Fig. 1A, B); antero- lateral corner rounded. Eye well developed, extremely elongate, projecting laterally, slightly less than 3 times as long as width of cornea; cornea functionally normal, much wider than eye- stalk; eyestalk narrow, broadened distally in distal half, without papilla (Fig. 1A, B). Antennular peduncle slender, 3-segment- ed. In male first segment longer than suc- ceeding 2 segments together, with tuft of setae on outer dorsal surface at distal end and distal fourth, 1 small spine on dorsal surface near distal end; second segment half as long as first, about 3 times as long as broad, with tuft of setae on outer dorsal sur- face at distal end, inner margin with 7 setae arranged at regular intervals; third segment shortest, less than 1.5 times as long as broad, armed with 4 setae arranged at reg- ular intervals on inner margin and with 2 setae at inner distal corner. Outer flagellum twice wider than inner, projecting antero- laterally and then anteriorly, with many fine setae on proximal part of inner margin (Fig. 1A). In female antennular peduncle more slender than that of male; second segment about 5 times as long as broad; third seg- ment about 3 times as long as broad. Outer VOLUME 111, NUMBER 4 Fig. 1. anterior end of female (allotype). flagellum narrower than inner, straight, without fine setae along inner margin (Fig. 1B). Antennal scale long, slender, extending to middle of second segment of antennular peduncle in male (Fig. 1A), extending slightly beyond anterior margin of first seg- ment of antennular peduncle in female (Fig. 1B), about 8.5 times as long as maximum Parapetalophthalmus suluensis, new genus, 851 oO gE —— SSS SS Sw, new species. A, anterior end of male (holotype); B, width at position of jointed spine marking end of naked part of outer margin; outer margin naked and concave in proximal %, setose in distal %; inner margin nearly Straight, setose except proximal third naked (Fig. 2A). Antennal peduncle slightly short- er than scale; first segment very short, pro- duced medially into triangular lobe; second segment with group of setae at distal sixth; 852 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Parapetalophthalmus suluensis, new genus, new species; A: allotype; B—G: holotype. A, antennal scale; B, mandible and mandibular palp; C, mandibles; D, maxillule; E, labrum; KE endopod of first thoracopod; G, endopod of second thoracopod. VOLUME 111, NUMBER 4 853 Fig. 3. Parapetalophthalmus suluensis, new genus, new species; A—F: holotype; G: allotype. A, endopod of third thoracopod; B, endopod of fifth thoracopod; C, distal part of endopod of fifth thoracopod; D, eighth thoracopod and penis; E, proximal part of fifth thoracopod; E proximal part of sixth thoracopod; G, endopod of seventh thoracopod. (END: endopod; EXP: exopod; P: penis). 854 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON —_——<—~_ LZ} TI} (es aa ey eee =e - —— —— a —— -— "EER — Y ————$——— 1) a Fig. 4. Parapetalophthalmus suluensis, new genus, new species; A—E: allotype; F—I: holotype. A, first ple- opod; B, second pleopod; C, third pleopod; D, fourth pleopod; E, fifth pleopod; E first pleopod; G, fifth pleopod; H, uropod and telson; I, distal part of telson. VOLUME 111, NUMBER 4 third segment equal to second in length, with group of setae at distal end. Female with flagellum 4-segmented, terminating in 2 short setae; male with flagellum about 18- segmented (Fig. 2A). Mandible of right side with lacinia mob- ilis (Fig. 2C). Palp large, slender, extending anteriorly beyond anterior margin of second segment of antennular peduncle in male and slightly beyond middle of second segment in female (Fig. 1B); first segment short; second segment cylindrical, nearly 7 times longer than broad, with 4 spine-like setae along outer margin and 7 along inner mar- gin; third segment % length of second, with flexure between second and third segments, armed with 9 strong setae, 3 on distal end, 2 on outer margin and 4 on inner margin, one of distal setae shorter than others, es- pecially in female (Figs. 1B, 2B). Maxilla and labrum as in Fig. 2D, E, respectively. Endopod of first thoracopod short and ro- bust, with epipod; inner lobe on ischium de- veloped, extending anteriorly to middle of merus, armed with several spine-like setae; carpopropodus longer than preceding 2 seg- ments combined, inner margin slightly con- vex, with group of setae on middle; claw fused with dactylus, long, inwardly curved (Fig. 2F); exopod wanting. Endopod of sec- ond thoracopod robust; lobe on ischium very large, quadrangular, overreaching distal mar- gin of merus, 4 times as long as broad, with about 27 long and short spine-like setae on inner and apical margins; carpopropodus el- liptical, shorter than preceding segment; claw fused with dactylus, robust, inwardly curved (Fig. 2G); exopod wanting. Endopods of third to fifth thoracopods extremely slender; carpopropodus as long as merus, undivided, with 5 groups of setae arranged at regular intervals on distal fourth of inner margin, each group composed of 2 or 3 setae; dactylus short, terminating in 2 short setae, without claw (Fig. 3A—C, E). Endopods of sixth to eighth thoracopods re- duced to 1- or 2-segmented, small rudi- ments with several setae (Fig. 3D, E G). Penis elongate, gradually thicker distally, 855 ending in hemispherical apex, without setae (Fig. 3D). Flagelliform part of thoracic ex- opods 9-segmented in third and fourth pairs and 10-segmented in fifth to eighth; basal plate narrow, with outer distal corner rounded (Fig. 3D). Abdominal somites increasing in length posteriorly, their length ratios are 1:1:1.4: 1.5:1.6:2.5; last abdominal somite more than 3 times as long as maximum breadth at posterior end; anterior 5 somites with pair of small blunt projections on posterior end of lateral margin. Pleopods of male biramous, well devel- oped; first pleopod with unsegmented en- dopod and 11-segmented exopod (Fig. 4F); second to fifth pleopods similar to each oth- er, without modified setae; exopod 12-seg- mented; endopod 6- or 7-segmented, slight- ly shorter than exopod; first segment long, with 2 groups of setae arranged transverse- ly, without definite pseudobranchial lobe (Fig. 4G). Pleopods of female biramous, in- creasing in length posteriorly; exopod very slender, not segmented or segmented indis- tinctly, with 2 to 6 setae on outer margin and 1 or 2 setae on apex; endopod fused with sympod, shorter than exopod in first to third pleopods, subequal in fourth and slightly longer in fifth (Fig. 4A—E). Exopod of uropod 2-segmented. Proxi- mal segment more than 4 times as long as broad, with both margins parallel; outer margin very slightly convex, naked except for distal end armed with 1 seta and 2 spines, proximal spine very small; inner margin very slightly concave, setose. Distal segment twice as long as broad at base, 0.28 of proximal segment in length, whole margin setose (Fig. 4H). Endopod of uro- pod unsegmented, extending to middle of distal segment of exopod, gradually nar- rowing distally (Fig. 4H). Telson rectangular, more than twice as long as broad, 0.7 of last abdominal somite in length. Lateral margins parallel, with 5 small, slender, equal-sized spines on each side. Distal margin armed with 9 grouped spines, each group consisting of 3 spines 856 with middle one usually longest; spines of outermost groups longer, barbed, jointed at base, 8 plumose setae present between 9 spine groups (Fig. 4H, I). Etymology.—The name suluensis is de- rived from the locality where the specimens were collected; it is an adjective agreeing in gender with the generic name. Remarks.—As previously discussed in the remarks section of the genus, the new species is easily distinguished from two species of the most closely related genus Pseudopetalophthalmus, P. japonicus Bra- vo & Murano, 1997, and P. australis (Pan- ampunnayil, 1982). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Parapetalophthalmus suluensis was col- lected in the depth of 285-306 m which was deeper than habitats of two species of Pseudopetalophthalmus, 74—130 m deep in P. japonius and subsurface in P. australis. Literature Cited Bravo, M. R., & M. Murano. 1997. Pseudopetalo- phthalmus japonius, new genus, new species (Mysidacea: Petalophthalmidae).—Journal of Crustacean Biology 17(4):725-—732. Panampunnayil, S. U. 1982. Description of a new spe- cies of Petalophthalmus (Mysidacea) with a re- vised definition of the genus.—Journal of Plankton Research 4(3):643—650. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):857—874. 1998. Diastylis tongoyensis, a new diastylid (Crustacea: Cumacea) from the northern central coast of Chile, with an amendment to the description of Diastylis crenellata Watling & McCann, 1997 Sarah Gerken and Les Watling Darling Marine Center, School of Marine Sciences, University of Maine, Walpole, Maine 04573 U.S.A. Abstract.—A new species, Diastylis tongoyensis, is described based on spec- imens from north central Chile. Diastylis tongoyensis is distinguished from other members of the genus by the unique combination of an opercular max- illiped 3, a short telson, and an unornamented carapace. Pereopods 3 & 4 of D. crenellata Watling & McCann, 1997 are also illustrated. Both belong to a group of 9 species in the genus Diastylis having an opercular maxilliped 3, reduced exopods on pereopods 3 & 4 of the female and a telson shorter than the uropod peduncles. Only two species belonging to Diastylis Say, 1818 are currently known from Chile, Diastylis argentata Calman, 1912 and D. gayi Nicolet, 1849. Diastylis argentata is described from off southern Chile, and D. gayi is described from Isla Grande de Chi- loé, also in southern Chile. Both sites are within the Magellanic biogeographic prov- ince, and D. gayi has not been reported sub- sequent to its description by Nicolet (1849). Diastylis tongoyensis is the first species from the Diastylidae described from the Peru-Chilean province (Fig. 1), which en- compasses the west coast of South America from about 40°S to 5°S. Methods Samples were collected with a small ben- thic dredge equipped with a net bag having a mesh opening of 425 wm, dragged by a small boat for 15-30 minutes in 3—30 me- ters of water. Because of the need to use the Specimens for genetic analysis, samples Were preserved directly in 95% EtOH. Drawings were prepared using a camera lu- cida on a Wild compound microscope. Body length was measured from the tip of the pseudorostral lobes to the posterior bor- der of the last somite. Family Diastylidae Bate, 1856 Diastylis Say, 1818 Diastylis tongoyensis, new species Figs. 2-11 Type material.—Holotype (USNM 260767), ovigerous female; paratypes (USNM 260768), 1 adult 3, 8 subadult 6d, 9 subadult 2 2, 15 manca 1; paratypes (Mu- seo Nacional de Historia Natural de Chile, Santiago CUM-no. 11153), 1 ovigerous 2, 7 subadult 22, 8 subadult ¢, 4 manca 1; Bahia Tongoy, Chile, 30°16.67’S, 71°30.25'W. Other material.—Bahia Tongoy, Chile, 30°16.67'S, 71°30.25'W, 4 ovigerous @ ¢, 30 subadult 2 2, 2 adult 6, 31 subadult 6, 132 manca 1, 381 individuals not sexed. Bahia Guanaquero, Chile, 30°11.66’S, 71°25.30'W, 6 subadult 2 2, 14 subadult d, 80 manca 1. Diagnosis.—Within the genus, distin- guished by the combination of unorna- mented carapace, telson considerably short- er than uropod peduncles, opercular max- illiped 3, pereonite 1 visible only dorsally, 858 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 70°W 30°S ee 0 — Bahia Tongoy Ro . ¢ = SANTIAGO 40°S— Fig. 1. Map of Chile, with enlargement of Bahia Tongoy area. VOLUME 111, NUMBER 4 859 Fig. 2. and rudimentary exopods on pereopods 3 & 4 in female. Description.—Ovigerous female, 6.0 mm. Pseudorostral lobes 0.2 carapace length, eyelobe 0.1 carapace length, with 3 lenses; antennal notch oblique; anteroven- tral corner slightly produced; ventral mar- gin lined with small teeth, otherwise cara- pace smooth. Branchial siphon not extend- ing beyond pseudorostral lobes. Pereonite 1 nearly obscured by carapace, slightly ex- posed dorsally. Pereonite 5 produced pos- teriorly as small acute lappet (Fig. 2A). Antennule elongate and stout, extending beyond pseudorostral lobes; peduncle arti- Diastylis tongoyensis, new species. A, adult 2; B, adult 6. cles 1 and 2 stout, article 3 slender; article 1 with 2 plumose setae distally; article 2 with 2 pedunculate setae distally; accessory flagellum of 2 articles, extending beyond article 1 of main flagellum; main flagellum of 4 articles, bearing 2 stout annulate setae (Fig. 3D). Antenna rudimentary; article 1 stout, 1 plumose seta distally; article 2 reduced, 1 simple seta distally (Fig. 3E). Mandible navicular with row of 11 lifting setae. Right lacinia mobilis very stout with two cusps, left lacinia mobilis slender with single cusp (Fig. 4A). Maxillule with 2 endites; outer endite with 860 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON o7 oi 2> . LL ZEEE S4 wa - Cara a Derren on sntnettlle oe SN esi WN Fig. 3. Diastylis tongoyensis, new species. Adult 2; A, maxilliped 1; B, maxilliped 2; C, maxilliped 3; D, antennule; E, antenna. VOLUME 111, NUMBER 4 C 861 Fig. 4. Diastylis tongoyensis, new species. Adult 2; A, mandibles; B, maxillule; C, maxilla. double row of stout simple setae distally, 1 slender simple seta on lateral margin, anterior face with several groups of very fine simple setae; inner endite with 5 setae terminally, 2 stout microserrate setae, 1 stout simple seta, 2 slender simple setae, anterior face with many fine simple setae; palp with 2 long stout simple setae distally (Fig. 4B). Maxilla with 3 endites; broad endite dis- tal half of medial margin with double row of fine simple setae, 5 large stout papose setae, proximal half of medial margin with row of 29 simple setae, 2 stout simple setae set below row, many groups of fine simple setae on anterior-medial face; inner narrow endite with 3 setae terminally, 2 stout sim- 862 ple setae, 1 broad blade-like seta; outer nar- row endite with 4 long setae terminally, 2 microserrate and 2 simple, lateral margin with 1 simple seta (Fig. 4C). Maxilliped 1 basis produced distally as blunt lobe, 2 hooks medially, medial margin beset with stout plumose setae; ischium partially fused; merus with 2 stout plumose setae; carpus and propodus subequal, both with field of slender plumose setae on me- dial face, propodus also bearing 2 long plu- mose setae distally; dactyl reduced, bearing 2 short simple setae distally (Fig. 3A). Maxilliped 2 endite with 5 long stout an- nulate setae; basis margins lined with fine hairs, distal medial corner with 3 long plu- mose setae; ischium compressed, bearing no setae; merus, carpus, propodus subequal in length, merus distal lateral corner bears single long plumose seta; carpus medial margin bearing row of slender sparsely plu- mose setae; propodus with 1 long plumose seta proximally, row of slender simple setae on distal half of medial margin; dactyl with 3 simple setae, 1 stout and 2 slender (Fig. 3B). Maxilliped 3 basis expanded, twice as long as all other articles together, medial margin with row of stout plumose setae, lat- eral margin beset with fine hair setae, distal lateral corner with group of long plumose setae, slightly proximally 2 stout setae, 1 simple and 1 plumose, distal medial corner produced as 2 teeth with short slender sim- ple seta between; ischium broad, incised distally, with no setae; merus medial distal margin with 2 long plumose setae and 1 short slender plumose seta, medial proximal margin beset with fine hairs; carpus re- flexed, without setae; propodus bearing 2 long plumose setae on medial distal margin; dactyl medial margin with 3 short plumose setae, tip with simple setae, 2 stout and 4 slender; exopod basal article with 1 proxi- mal and 1 distal plumose seta, flagellum bearing many long plumose setae, basis and flagellum together half endopod basis length (Fig. 3C). Pereopod 1 basis as long as carpus, pro- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON podus, dactyl together, anterior and poste- rior margins with many long plumose setae, distal margin with row of long plumose se- tae on medial margin, lateral margin pro- duced as 2 large teeth, basis body with stout triangular scales that appear as teeth on margins; ischium with tooth on distal cor- ner, otherwise unarmed; merus longer than ischium, unarmed; carpus, propodus, dactyl subequal in length, carpus with few delicate setae, propodus distal corner with single long plumose seta; dactyl with 3 plumose setae distally; plumose setae on propodus and dactyl having closely set short setules, not similar to plumose setae on basis; ex- opod shorter than endopod basis, basal ar- ticle with 5 short plumose setae, flagellum with many long plumose setae (Fig. 5A). Pereopod 2 basis slightly expanded, as long as next 4 articles together, anterior and posterior margins with many long plumose setae; ischium short, posterior margin with 1 long plumose seta; merus 4 times ischium length, distal corner with 1 plumose seta; carpus longer than propodus and dactyl to- gether, unarmed; propodus half as long as dactyl, 1 simple seta distally; dactyl with several simple setae; exopod longer than endopod basis, exopod basal article shorter than flagellum with 3 slender plumose setae posteriorly, flagellum with many long plu- mose setae (Fig. 6B). Pereopod 3 basis as long as remaining articles combined, with 2 pedunculate setae at distal corner, armed with several long plumose setae and 2 smaller annulated se- tae; ischium margin with several plumose setae and several annulate setae; merus as long as carpus and propodus together, distal corner with single annulate seta, lateral face with 1 annulate seta; carpus margin lined with long annulate setae, carpus as long as dactyl and propodus together; propodus un- armed; dactyl 0.5 width of propodus, single very stout simple seta distally; exopod re- duced, 2 small articles, distal article bearing single annulate seta and several fine setae (Fig. 7B). Pereopod 4 basis as long as merus and VOLUME 111, NUMBER 4 ? Fig. 5. carpus together, posterior margin with sev- eral long stout plumose setae, 1 pedunculate seta and 3 slender annulate setae on outside; ischium with 5 long annulate setae; merus half basis length, posterior margin with 4 long annulate setae; carpus with row of long stout annulate setae distally, few slen- 863 Diastylis tongoyensis, new species. A, adult 2 pereopod 1; B, adult d pereopod 1. der annulate setae not on margin; propodus with 1 extremely stout annulate seta distal- ly; dactyl reduced, 0.5 width of propodus, bearing single short stout seta distally; ex- opod reduced, 2 articles, distal article with 1 plumose seta (Fig. 6A). Pereopod 5 basis as long as merus and 864 Fig. 6. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON _ ‘Ss 0.25 mm Diastylis tongoyensis, new species. Adult 2; A, pereopod 4; B, pereopod 2. 865 VOLUME 111, NUMBER 4 HASSE 0.25 mm 7. Diastylis tongoyensis, new species. A, adult d pereopod 4; B, adult 2 pereopod 3. 866 carpus together, distal corner with 1 plu- mose seta, proximally 2 plumose setae, dis- tally 1 stout annulate seta; ischium distal corner with single annulate seta, row of 3 annulate setae laterally; merus *% basis length, margins with few slender annulate setae; carpus with 2 long annulate setae dis- tally, 5 short annulate setae on outside; pro- podus with single extremely stout annulate seta distally; dactyl reduced, 0.5 width of propodus, with 1 short stout simple seta dis- tally (Fig. 8D). Telson length 0.7 length of uropodal pe- duncles, pre-anal and post-anal sections subequal in length, post-anal portion very slender with 3 pairs of lateral setae, 1 pair of short stout simple setae terminally (Fig. 9A). Uropod peduncles longer than rami, -pe- duncle medial margins armed with 9 or 10 stout setae. Uropod endopod of 3 articles; article 1 medial margin with 4 stout setae, lateral margin with 2 pedunculate setae, lat- eral distal corner with single seta; article 2 medial margin with 3 setae, lateral distal corner with single seta; article 3 medial margin with 2 setae, lateral margin with single pedunculate seta, | large stout seta and 1 small slender seta on distal margin. Uropod exopod of 2 articles; article 1 one- fourth length article 2, armed with 1 short seta; article 2 lateral margin lined with sim- ple setae, medial distal corner with single seta, distal margin with 2 long stout setae. All setae on medial margins, both peduncle and endopod, and large seta on the endopod distal margin, short, stout setae with single setule sub-terminally (Fig. 9A). Adult male, 5.8 mm. Carapace pseudo- rostral lobes 0.2 total carapace length, not upturned; antennal notch completely oblique; anteroventral corner slightly ser- rate, otherwise smooth. Carapace generally smooth. Eye large, 0.7 length of pseudo- rostral lobes, with at least 3 lenses. Bran- chial siphon not extending beyond pseu- dorostral lobes. Pereonite 1 almost com- pletely covered by carapace, exposed dor- sally. Pereonite 5 produced acutely PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON posteriorly, much longer than in female, with 4 long terminal setae on lappet. Pleon- ites 1—5 with 2 setae on each posterior-ven- tral corner (Fig. 2B). Antennules much longer than pseudoros- tral lobes, peduncle articles very stout; ar- ticle 1 equal to articles 2 and 3 together, margins beset with fine hairs, 2 simple short setae distally; articles 2 and 3 subequal, ar- ticle 2 unarmed; article 3 entire distal mar- gin bearing very slender setae, equal in length to peduncle of main flagellum; ac- cessory flagellum of 3 articles, proximal ar- ticle bears single pedunculate seta proxi- mally; main flagellum of 5 articles, distal article bears 2 simple setae, as long as setae on article 3 (Fig. 10A). Antennal peduncle of 5 articles; first 4 articles equal in length to article 5, articles 1 and 2 subequal; article 1 with 1 plumose seta; article 2 with 2 plumose setae; articles 3 and 4 subequal, together equal to article 2, both unarmed; article 5 stout, with 19 rows of fine setae on anterior margin; fla- gellum extends at least to end of uropod rami, comprising at least 20 articles, each article bearing several rows of fine setae, articles increasing in length and decreasing in width distally (Fig. 8A). Maxilliped 3 basis expanded, medial margin with many short plumose setae; is- chium expanded, incised distally, unarmed; merus medial margin with 2 short plumose setae; carpus reflexed, unarmed; propodus medial margin with 2 long simple setae, distally 2 short simple setae; dactyl margins with several simple setae, 2 long simple se- tae distally. Exopod 0.7 endopod basis; ex- opod basal article expanded, unarmed; fla- gellum bearing long plumose setae (Fig. 11B). Pereopod 1 basis equal to all other arti- cles together, with plumose setae on ante- rior, posterior, and distal margins, increas- ing in length distally, medial margin with several teeth; ischium shorter than merus, unarmed; merus with 2 small setae distally; carpus with 2 small simple setae on margin; propodus longer than carpus, with 3 slender VOLUME 111, NUMBER 4 867 0.25 mm VN AS 0 Fig. 8. Diastylis tongoyensis, new species. A-C, adult d, D, adult 2; A, antenna; B, pereopod 5; C, pleopod 2; D, pereopod 5. 868 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A ZO SSA GAS SS LAVE-—WUVE iD VOLUME 111, NUMBER 4 895 Fig. 2. Lamoha hystrix, new species. Holotype female, carapace width 38.7 mm, carapace length 48.9 mm, BPBM 511810. A, carapace, dorsal view; B, abdomen. SSS Fig. 3. Lamoha hystrix, new species. Holotype female, carapace width 38.7 mm, carapace length 48.9 mm, BPBM 511810. a, right merus of fifth ambulatory leg; b, subchelate structure on right fifth ambulatory leg. Scales = 5.0 mm. 896 angular (length to width ratio 1.26); rostrum bifurcated, with relatively elongate base; posterolateral margin with row of granules; protogastric region with 4 spines; mesogas- tric region with 1 spine; metagastric region granulated; subhepatic region with 2 lateral spines and 5 other spines on surface; P2—4 elongate, M2-—4 relatively wide, with 14— 18, 19-20, 18 dorsal spines respectively; P5 with 3 dorsal spines and 8—9 ventral spines; Pr4 with 6—7 spines; subchelate margin of propodus of P5 with 9—10 spines. Description of holotype female.—Cara- pace longitudinally rectangular, regions well defined (Figs. 1A, 2A). Entire carapace and pereiopod surfaces with short, stiff, simple setae. Rostrum well developed, base relatively elongate, distal part bifurcated with 2 sharp spines. Pseudorostral spines as long as rostral spines, sharp. Supraorbital spines very long (Figs. 1A, 2A). Anterolat- eral margin with 2 lateral and 2 subdorsal spines. Posterolateral margin gently con- vex, lined with a row of small sharp and rounded granules along “linea homolica”’ (Fig. 2A). Protogastric region with 3 large and | small (inner posterior) spines. Me- sogastric region with | short, sharp spine. Metagastric region granulated but without spines. Branchial regions covered with fine granules. Subhepatic region with 2 lateral spines, 1 small submarginal spine, 1 sub- dorsal spine and 3 subventral spines (Figs. 1B, 2A). Posterior part of pterygostomial region covered with granules and spinules. Gastro-cervical groove deep, contiguous medially. Branchio-cardiac groove deep (Fig. 2A). Basal antennal article with 1 in- ner spine. Antennular peduncle with 1 sharp spine. Posterior carapace margin sinuous, median part concave. Main proepistomal spine long, sharp; lateral spines distinctly shorter (Fig. 1B). Merus of third maxilliped with group of 2—3 spines medially, rest of margin with scattered spines and spinules; ischium with outer margin anterior % of outer margin spinate, with distinct granu- lated longitudinal median ridge; exopod slender, reaching to mid-point of outer mar- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gin of merus, proximal half with granule- lined longitudinal median ridge. P1 (chelipeds) subequal, elongate, slen- der (Fig. 1A). Carpus with 3—4 sharp, curved spines on inner distal margin; outer surfaces of chelae with several rows of spi- nules, spines and granules, those along out- er surface flatter or blunter, without any trace of pigmented spot (Fig. 1C). Fingers elongate, distal part curving inwards, dac- tylus with strong longitudinal ridge and sul- cus, distal one-third of dactylus and cutting margins pigmented dark brown in preser- vative, cutting edges smooth, blade-like (Fig. 1C). P2 and 3 longest (Fig. 1A). Rest of armature on pereiopods as follows: M5 with 3 dorsal and single row of 8—9 ventral spines (Fig. 3a); M4 with 18 dorsal and 2 rows of ventral spines (posterior row with 17-18 spines, anterior row with 21 spines); M3 with 19—20 dorsal spines and 2 rows of ventral spines (posterior row with 19—23 spines, anterior row with 28 spines); M2 with 14-18 dorsal spines and 2 rows of ventral spines (posterior row with 16—21 spines, anterior row with 19-26 spines); M1 with 14 dorsal spines (with 1 distally positioned) and 2 rows of ventral spines (posterior row with 12—13 spines, anterior row with 15-17 spines); Pr4 with 6—7 spines of which 2 are distally positioned, movable and bracketing base of dactylus; D3 and D4 with 19 spines on ventral mar- gin; D2 with 16—18 spines on ventral mar- gin. Dorsal armature of all pereiopods with median spines largest; spines on ventral ar- mature of pereiopods progressively larger towards distal end, with numerous granules of varying sizes at proximal surface which makes counting of exact number of spines difficult; rows of spines on ventral margins not linear but uneven, especially along proximal part; all spines curving outwards. Dactylus and distal part of propodus of P5 forming subchelate structure; anterior tip of propodus with 4—5 movable spines; sub- chelate margin of propodus with 9-10 spines; ventral margin of dactylus with 4 movable spines (Fig. 3b). Basis-ischium of VOLUME 111, NUMBER 4 Pl with 1 sharp dorsal spine and several smaller spines and granules; those of P2—4 with 2 dorsal spines bracketing merus; that of P5 with 1 median dorsal spine. Coxa of P1 with 1 sharp on inner dorsal angle and 1 large granule on outer dorsal angle; those of P2—4 with 2 sharp spines on dorsal mar- gin bracketing basis-ischium; that on P5 without spines or spinules. Abdomen covering entire thoracic ster- num; telson triangular, with distal part of lateral margins concave (Fig. 2B). Remarks.—Lamoha hystrix, new species, appears to be closest to L. longipes (Alcock & Anderson, 1899) (Indian Ocean), L. mu- rotoensis (Sakai, 1979) (Japan, Taiwan, and Madagascar and Seychelles with doubt) and L. inflata (Guinot & Richer de Forges, 1981) (Loyalty Islands, Samoa and Tua- motu). This group of species is essentially defined by the form of the frontal and su- praorbital margins, the proepistome pos- sessing a distinct vertical spine and the pro- podus of Pr5 being very short with the pro- podal finger of the subchelate process very low (Guinot & Richer de Forges, 1995: 444). In addition, all four species have sim- ilar fourth ambulatory meri which are armed with spines along the ventral margin. None of these species, however, have the dorsal margin of the merus armed with spines as in L. hystrix. The spines on the dorsal margin of the fourth leg in L. hystrix are relatively small but very distinct (Fig. 3a). The carapace features of L. hystrix ap- pear to be closest to L. longipes with re- gards to the posterolateral margin lined with small granules. In addition to the ear- lier mentioned presence of spines on the dorsal margin of the fourth ambulatory leg, L. hystrix, can easily be separated from L. longipes by the its proportionately longer ambulatory meri (M4 length to maximum width ratio 8.1 in L. hystrix, 7.5—7.6 in L. longipes), larger number of spines on the ambulatory meri (e.g., M4 with 18 in ZL. hystrix, 13-14 dorsal spines in L. longipes), presence of spines on the ventral margin of 897 Pr4 (absent in L. longipes) (cf. Alcock 1901: pl. 6 fig. 25), and the larger number of teeth on the subchelate margin of the propodus of P5 (9-10 spines in L. hystrix, 6 in L. longipes) (cf. Williams 1974:489, fig. 8). The outer surface of the female che- la also lacks a dark spot. A note on L. longipes is relevant here. The carapace proportions for the various re- ported specimens of L. longipes seem dif- ferent, although this may be a result of dif- ference in sizes. The specimen (apparently a male 39.4 by 30.6 mm, carapace length to width) figured by Alcock (1901: pl. 6 fig. 25) has a carapace length to width ratio of 1.29. Alcock (1901:69) also mentions a large ovigerous female measuring 38.0 by 30.0 mm (ratio 1.27). The female specimen of L. longipes from the Seychelles exam- ined and figured by Guinot & Richer de Forges (1995:447, fig. 56a, b), however, measures 33.2 by 29.0 mm, and has a length to width ratio of 1.14, that is, the carapace is more squarish. The spines on the protogastric and mesogastric regions of this specimen also appear to be lower and more scattered, with the rostral base very short, compared to the material described and illustrated by Alcock & Anderson (1899:6), Alcock (1899:15, pl. 1 fig. 1; 1901:69, pl. 6 fig. 25) and Doflein (1904: pl. 11 fig. 1, 2; pl. 50 fig. 4; pl. 51 fig. 1; pl. 52 figs. 1-5). Whether this material is really L. longipes cannot be ascertained as only one specimen from the Seychelles was obtained. In any case, Guinot & Richer de Forges (1995:447) had referred the speci- men to the species with reservations. The carapace proportions of L. hystrix, new spe- cies (length to width ratio 1.26) are com- parable with those of L. longipes sensu stricto. The positions and strengths of the spines on the gastric region of L. hystrix, new spe- cies, are very similar to that on L. muro- toensis, but L. murotoensis is easily sepa- rated by the strong longitudinal ridge on its posterolateral margin. Lamoha inflata is a 898 very distinctive species with the gastric car- apace regions unarmed. In addition to L. murotoensis and L. in- flata, three other species are known from the Pacific, i.e., L. williamsi (Takeda, 1980) (Kyushu-Palau Ridge), L. personata (Guin- ot & Richer de Forges, 1981) (Polynesia, Vanuatu, Samoa, Kiribati, Australia), and L. futuna (Guinot & Richer de Forges, 1995) (Wallis and Futuna Islands) (Guinot & Richer de Forges 1995). These species, however, differ markedly from L. hystrix in a multitude of pereiopod and carapace fea- tures (Guinot & Richer de Forges 1995). According to the collection data on the specimen of Lamoha hystrix, new species, it was caught in a trap baited with mullet set in the afternoon and left for 24 hours between 305 to 366 metres. The colour in life was recorded as “‘rose-pink’’. The ecol- ogy and habits of the species is not known, but is probably similar to that of other Pa- cific Lamoha species (Guinot et al. 1995). Etymology.—The species is named after the porcupine (Hystrix) because of its spiny carapace and legs. The name is used as a noun in apposition. Acknowledgments Thanks are due to Lu Eldredge for his kind help and hospitality during my stay in the Bernice P. Bishop Museum. I am also grateful to Daniéle Guinot (Paris Museum) for her useful comments with regards to the specimen. Mr. S. H. Tan and Mr. H. K. Yip kindly photographed the specimen. Literature Cited Agassiz, L. 1859. Remarks on fishes from Lake Nic- aragua.—Proceedings of the Boston Society of Natural History 6(1856—1859):407—408. Alcock, A. 1899. An account of the deep sea Brach- yura collected by the Royal Indian Marine Sur- vey Ship Investigator. Calcutta, 85 pp. . 1901. Catalogue of the Indian decapod Crus- tacea in the collection of the Indian Museum. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Part I. Brachyura. Fascicle I. Introduction and Dromides or Dromiacea (Brachyura Primigen- ia). Trustees of the India Museum, Calcutta, 80 pp:, pls. 1-8. , & A. R. Anderson. 1899. Natural history notes from H.M. Royal Indian marine survey ship ‘Investigator’, Commander T. H. Heming R. N., commanding. Series 3, number 2. An ac- count of the deep-sea Crustacea dredged during the surveying-season of 1897—98.—Annals and Magazine of Natural History (7)3:1—27, 278— 292. Doflein, F 1904. Brachyura. Wissenschaften Ergeben- isse Deutschen Teifsee ‘Valdivia’ Expedition 6: i-xiv, 1-314, figs. 1-68, pls. 1-57. Guinot, D., & B. Richer de Forges. 1981. Homolidae, rares ou nouveaux, de |’ Indo-Pacifique (Crus- tacea, Decapoda, Brachyura).—Bulletin du Mu- séum national d’Histoire Naturelle, Paris (4)3 section A(2):523—581, pls. 1-8, maps 1-2. ,& . 1995. Crustacea Decapoda Brach- yura: Révision de la famille des Homolidae de Haan, 1839. In: A. Crosnier, ed., Résultats des campagnes MUSORSTOM, vol. 13.—Mémo- ires du Muséum national d’Histoire Naturelle, Paris 163:283-517. , D. Doumenc, & C. C. Chintiroglou. 1995. A review of the carrying behaviour in brachyuran crabs, with additional information on the sym- bioses with sea anemones.—Raffles Bulletin of Zoology 43:377—416. Ng, P. K. L. 1998. Lamoha, a replacement name for Hypsophrys Wood Mason & Alcock, 1891 (Brachyura, Homolidae), a junior synonym of Hypsophrys Agassiz, 1859 (Pisces, Teleostei, Cichlidae).—Crustaceana 71(1):121—125. Sakai, T. 1979. Description of three new species of crabs of the family Homolidae from Japan.— Researches in Crustacea, Tokyo 9:1-8 (En- glish), 8-12 (Japanese), 1 color frontispiece. Takeda, M. 1980. A small collection of crabs from the Kyushu-Palau Submarine Ridge, with descrip- tion of a new species of the Homolidae.—Mi- cronesica 16(2):279—287. Williams, A. B. 1974. A new species of Hypsophrys (Decapoda: Homolidae) from the Straits of Florida, with notes on related crabs.—Proceed- ings of the Biological Society of Washington 87(42):485—492. Wood-Mason, J., & A. Alcock. 1891. Natural History Notes from H. M. Indian Marine Survey Steam- er ‘Investigator’, Commander R. E Hoskyn, R. N., commanding.—No. 21. On the Results of the last Season’s Deep-sea Dredging.—Annals and Magazine of Natural History (6)7(39):258— Diz: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):899—907. 1998. A new freshwater crab of the genus Neostrengeria Pretzmann, 1965, from Colombia (Crustacea: Decapoda: Brachyura: Pseudothelphusidae), with a Key to the species of the genus Martha R. Campos and Rafael Lemaitre (MRC) Universidad Nacional de Colombia, Instituto de Ciencias Naturales, Apartado Aéreo 53416, 114 Santa Fé de Bogota 2, Colombia, S.A.; (RL) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. Abstract.—A new species of freshwater crab of the pseudothelphusid genus Neostrengeria Pretzmann, 1965, N. perijaensis, is described and illustrated. This discovery means the genus now contains a total of 17 species and 2 subspecies, all endemic to the Eastern Andes of Colombia. The general char- acteristics of the genus are given, and a key for the identification of the species based primarily on the morphology of the male first gonopod, is presented. The genus Neostrengeria Pretzmann, 1965, of the family Pseudothelphusidae, comprises a group of freshwater crabs that live in mountain springs and streams on the slopes and high plains of the Eastern Andes of Colombia (about 3° to 9°40’N, 73° to 74°50’'W), at altitudes ranging from 400 to 3000 m above sea level. The systematics of this genus were clarified by Rodriguez (1982), and more recently were reviewed by Campos (1992, 1994). The geographical distribution of the genus was discussed by Campos & Rodriguez (1985) and Campos (1992, 1994). The new species described herein, N. perijaensis, was collected from humid habitats south of the Serrania of Per- ya, at altitudes ranging between 1200 and 1800 m above sea level. With this discov- ery, the genus now contains 17 species and two subspecies. The general carapace morphology of most species of Neostrengeria is very sim- ilar. They are distinguished primarily by the relative length of the exognath of the third maxilliped, which is 0.5 to 0.7 times as long as the ischium; the orifice of the efferent branchial channel, which is open; and by the male first gonopod which has a distinct lateral lobe usually divided in two halves, forming an accessory lobe. The shape of the apex of the first gonopod varies accord- ing to species, and is either oval, oblong, or expanded. A key to the species and subspe- cies of the genus is presented, based almost exclusively on the morphology of the first gonopod. The terminology used for the morphology of the first gonopod follows Smalley (1964) and Rodriguez (1982). The material is deposited in Museo de Historia Natural, Instituto de Ciencias Na- turales, Universidad Nacional de Colombia, Santa Fé de Bogota (ICN-MHN); National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM); and Instituto Venezolano de Investigaciones Cientificas (IVIC). The abbreviations cb and cl indicate carapace breadth and length, respectively. Color nomenclature used fol- lows Smithe (1975). Coordinates of col- lecting sites were taken using a Geograph- ical Positioning System (GPS). Family Pseudothelphusidae Rathbun, 1893 Tribe Strengerianini Rodriguez, 1982 Genus Neostrengeria Pretzmann, 1965 Neostrengeria perijaensis, new species Riese Holotype.—Quebrada El Zumbador, Ver- eda El Zumbador, Corregimiento La Vic- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Neostrengeria perijaensis, new species, male paratype, cb 17.5 mm, cl 11.0 mm (USNM 276145). A, dorsal view; B, ventral view. VOLUME 111, NUMBER 4 901 FE Fig. 2. Neostrengeria perijaensis, new species, male holotype, cb 22.0 mm, cl 13.5 mm (ICN-MHN-CR 1561). A, left first gonopod, caudal view; B, same, lateral view; C, same, cephalic view; D, same, mesial view; E, same, apex, distal view; E left second gonopod, caudal view; G, left third maxilliped, external view; H, opening of left efferent branchial channel, external view. Scales equal 2 mm (A-E, G), and 1 mm (EF 4H). 902 toria de San Isidro, Municipio La Jagua de Ibirico, Serrania de Perijaé, Cesar Depart- ment, Colombia, 9°33’10.8”N, 73°9'10.7’"W, 1300 m alt., 13 Mar 1996, leg. M. R. Cam- pos: 1 d, cb 22.0 mm, cl 13.5 mm, (ICN- MHN-CR 1561). Paratypes.—Same locality data as holo- iyo I SG, Go NS mn, el) LilO iim, (USNM 276145), 1 3, cb 15.0 mm, cl 9.3 mm, 4 &, cb range 20.7—-11.6 mm, cl range 12.9-10.0 mm (ICN-MHN-CR 1562). Non-paratypes.—Quebrada El Zumba- dor, Vereda El Zumbador, Corregimiento La Victoria de San Isidro, Municipio La Jagua de Ibirico, Serrania de Perija, Cesar Depart- ment, Colombia, 9°33'14.2”N, 73°9'17.8"W, 1270 m alt., 15 Mar 1996, leg. M. R. Cam- joe 6.5 Go W653 ls mn Cl iO4. 72 mm, 5 2, cb range 19.5-14.4 mm, cl range 12.0—8.8 mm (ICN-MHN-CR 1565).—Alto de Cantarrana, Vereda El Zumbador, Cor- regimiento La Victoria de San Isidro, Mun- icipio La Jagua de Ibirico, Serrania de Per- ja, Cesar Department, Colombia, 9°32'49.8"N, 73°8'46.9"W, 1800 m alt., 18 Mar 1996, leg. M. R. Campos: 1 6, cb 17.1 mm, cl 10.6 mm (IVIC), 11 3, cb range 18.4—11.1 mm, cl range 11.2—7.4 mm, 10 2, cb range 23.2-13.0 mm, cl range 13.8— 8.5 mm (ICN-MHN-CR 1568).—Finca Per- alonso, Vereda El Zumbador, Corregimien- to La Victoria de San Isidro, Municipio La Jagua de Ibirico, Serrania de Perija, Cesar Department, Colombia, 9°32'37.8'N, 73°9'42.5"W, 1320 m alt., 19 Mar 1996, leg. M. R. Campos: 6 ¢, cb range 20.1—12.8 mm, cl range 12.2-8.1 mm, 11 2, cb range 21.2-13.7 mm, cl range 13.3-8.4 mm (ICN-MHN-CR 1569). Type locality.—Quebrada El Zumbador, Vereda El Zumbador, Corregimiento La Victoria de San Isidro, Municipio La Jagua de Ibirico, Serrania de Perija, Cesar Depart- ment, Colombia, 9°33'10.8”N, 73°9'10.7’"W, 1300 m alt. Diagnosis.—First male gonopod straight, wide in caudal view; apex expanded ce- phalically into acuminate projection form- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ing strong, wide-based spine; mesial lobe developed into distinct elongate projection. Description of holotype.—Carapace (Fig. 1A) with cervical groove straight, shallow, ending some distance from lateral margin. Anterolateral margin with shallow depres- sion behind external orbital angle, followed by series of papillae on anterior half; pos- terior half smooth. Postfrontal lobes small, rounded, delimited anteriorly by 2 depres- sions; median groove shallow. Surface of carapace in front of postfrontal lobes in- clined anteriorly, depressed towards mid- line. Front rounded, lacking distinct upper border in frontal view, slightly bilobed in dorsal view; lower margin strongly sinuous in frontal view. Orbital margins each with row of small tubercles. Dorsal surface of carapace smooth, covered by small papil- lae; regions distinctly marked. Third max- illiped with merus having sharp angle on distal half of external margin; exognath ap- proximately 0.58 times length of ischium (Fig. 2G). Orifice of efferent branchial channel irregularly ovate (Fig. 2H). First pereiopods heterochelous; left che- liped larger than the right. Merus with 3 longitudinal crests as follows: upper one with rows of tubercles, internal lower one with rows of teeth, and external lower one with few tubercles. Carpus with 5 tubercles on internal crest and prominent blunt spine distally. Palms of both chelipeds smooth, swollen. Fingers of chelae not gaping when closed, tips crossing; outer and inner sur- faces with rows of small tubercles. Walking legs (pereiopods 2—5) slender (Fig. 1A). Dactyli elongated, each about 1.6 times as long as propodi, with papillae and 5 longitudinal rows of large spines dimin- ishing in size proximally. Spines and papil- lae on each dactylus arranged as follows: 1 anterolateral row and 1 anteroventral row each with 5 spines and 2 intercalated pa- pillae; 1 external row with 4 spines, 3 in- tercalated papillae and 1 pair of proximal papillae; and 1 posteroventral row and 1 posterolateral row each with 4 spines. First gonopod (Fig. 2A—E) straight, wide; VOLUME 111, NUMBER 4 mesial border straight on subdistal portion; with wide, deep notch in caudal view. Ac- cessory lobe rounded distally, slightly shorter than lateral lobe and directed cau- dally; lateral lobe wide, distal end rounded, directed cephalically, with prominent spine- like process on external margin (Fig. 2A— C). Apex in distal view (Fig. 2E) expanded cephalically into acuminate projection forming strong, wide-based spine; surface covered by dark colored spines; mesial lobe developed into distinct elongated projec- tion, surface rough; mesocaudal projection of spermatic channel terminating acutely, perpendicular to apical expansion (Fig. 2D, E). Second gonopod (Fig. 2F) with spinules distally; tip cup-shaped. Color.—The alcohol preserved holotype is dark brown (near 223, Raw Umber), with pale brown (Verona Brown, 223 B) specks on the dorsal side of the carapace. The walking legs are brown (Mars Brown, 223 A) dorsally, and buffy-brown (Sayal Brown, 223C) ventrally. The large chela is pale brown (Verona Brown, 223C) dorsally, and buffy-brown (Sayal Brown, 223C) ven- trally. The small chela is reddish-brown (Tawny, 38). The ventral surface of the car- apace is buffy-brown (Sayal Brown, 223C). Habitat.—The specimens were collected in shaded, moist banks of springs and small streams. They were found in soft mud, un- der rocks, or in burrows. The largest pop- ulations were found in one location at Alto de Cantarrana, Vereda El Zumbador, Cor- regimiento La Victoria de San Isidro, Mun- icipio La Jagua de Ibirico, Cesar Depart- ment. Etymology.—tThe specific name refers to the Serrania de Perija, where all the speci- mens were collected. Remarks.—This species is most similar to Neostrengeria lobulata Campos, 1992. The two can be differentiated by features of the first gonopod. The first gonopod (in caudal view) of N. lobulata is narrow and constricted near its midsection, while the first gonopod of N. perijaensis is wide and straight (cf. Campos 1992). The mesial lobe 903 of the first gonopod of N. perijaensis is de- veloped into a distinct elongate projection, while this lobe is rudimentary, represented by only a subterminal swelling in N. lobu- lata. The apex of the first gonopod of N. lobulata is formed by a wide, irregularly shaped expansion with curved borders that is projected cephalically into an acute lobe, while in N. perijaensis the apex consists of an acuminate projection ending in a strong, broad-based spine. Key to the species of the genus Neostrengeria based primarily on the male first gonopod 1. Laterodistal expansion of first gonopod cu.ved, forming wide lobe (Fig. 3A) N. botti Rodriguez & Tiirkay, 1978 — Laterodistal expansion of first gonopod not curved, lacking lobe 2. Mesial lobe of first gonopod triangular 3 — Mesial lobe of first gonopod semicir- Cuan (BISA yy watts hh i ka N. guenteri Pretzmann, 1965 3. First gonopod with lateral and acces- sory lobes distinctly separated — First gonopod with lateral and acces- sory lobes not separated from gonopod 12 4. Apex of first gonopod with outer sur- LACE SIMO OWN IA, Sia Meloke Get eee eele Oe 5) — Apex of first gonopod with outer sur- face SpinulOse™. Jaan ws ceed se 17 5. Accessory lobe of first gonopod sube- qual in length to lateral lobe ........ 6 — Accessory lobe of first gonopod dis- tinctly shorter than lateral lobe 6. Accessory lobe of first gonopod un- armed (Fig. 3C) WN. gilberti Campos, 1992 — Accessory lobe of first gonopod armed with spines (Fig. 3D, E) N. aspera Campos, 1992 7. Margins of accessory lobe of first gon- opod smooth in caudal view — Margins of accessory lobe of first gon- opod festooned in caudal view (Fig. 3F) Ses N. monterrodendoensis Bott, 1967 8. Longitudinal rows of spines on dactyli of walking legs each with 3 to5 spines 9 — Longitudinal rows of spines on dactyli of walking legs each with 6 to 10 spines 904 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON CPi non Fig. 3. Left first gonopod of species of Neostrengeria Pretzmann, 1965: A-E, caudal view; G, apex, distal view. A, N. botti Rodriguez & Tiirkay, 1978; B, N. guenteri Pretzmann, 1965; C, N. gilberti Campos, 1992; D, N. aspera Campos, 1992; E, accessory lobe of same; E N. monterrodendoensis Bott, 1967; G, N. lindigiana (Rathbun, 1897). 1, laterocaudal expansion; 2, mesial lobe; 3, lateral lobe; 4, accessory lobe; 5, mesocaudal projection of spermatic channel. Scales equal 1 mm. VOLUME 111, NUMBER 4 905 CPinten Fig. 4. Left first gonopod of species of Neostrengeria Pretzmann, 1965, caudal view. A, N. charalensis Campos & Rodriguez, 1985; B, N. macarenae Campos, 1992; C, N. lobulata Campos, 1992; D, N. libradensis libradensis Rodriguez, 1980; E, N. 1. appressa Campos, 1992. Scales equal 1 mm. 906 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON OOP inz4 E Fig. 5. Apices of left first gonopod of species of Neostrengeria Pretzmann, 1965, distal view. A, N. tonensis Campos, 1992; B, N. tencalanensis Campos, 1992; C, N. boyacensis Rodriguez, 1980; D, N. lasallei Rodriguez, 1980; E, N. niceforoi (Schmitt, 1969). 1, cephalic margin; 2, laterocaudal margin. Scales equal 1 mm. 9. Apex of first gonopod oval in distal 11. Lateral lobe of first gonopod short, dis- VIEW ris aia o Rese ae cat Ganapati 11 tal margin semicircular in caudal view — Apex of first gonopod semicircular in (Hig. AAYS fs he yr ee N. charalensis distal view, expanded cephalically (Fig. Campos & Rodriguez, 1985 BG) eae 62 N. lindigiana (Rathbun, 1897) — Lateral lobe of first gonopod long, dis- tal margin broadly rounded (Fig. 4B) suse eRELS N. macarenae Campos, 1992 10. Walking legs unusually long, total length (measured from coxa to tip of dactyl) of each about 1.3 times cara- pace width .... WN. sketi Rodriguez, 1985 — Walking legs normal in length, total 12. Laterodistal margin of first gonopod widening distally, forming lobe (Fig. length (measured from coxa to tip of 4C) Hewes a lobulata Campos, 1992 dactyl) of each subequal to carapace Se Laterodistal Margin of first gonopod not Wilting earshot ae bate ter ce oo widening distally, not forming lobe .. 13 ... N. macropa (H. Milne Edwards 1853) 13. Mesocaudal projection of spermatic VOLUME 111, NUMBER 4 channel of first gonopod terminating in ACICEOIICEPLO;ECHON aan. ss 4. 14 — Mesocaudal projection of spermatic channel of first gonopod terminating in acute simple projection (Fig. 2E) 35 eee eee N. perijaensis, new species 14. Lateral lobe of first gonopod directed PIC AlllWar mn tee eer a Ae eee a Se eee 15 — Lateral lobe of first gonopod curved ce- phalically 15. Distal portion of accessory and lateral lobes of first gonopod separated by deep notch in lateral view (Fig. 4D) N. libradensis libradensis Rodriguez, 1980 — Distal portion of accessory and lateral lobes of first gonopod not separated by deep notch, nearly continuous, in lateral view (Fig. 4E) N. libradensis appressa Campos, 1992 16. Cephalic margin of apex of first gono- pod constricted subdistally, terminating in spine (Fig. 5A) N. tonensis Campos, 1992 — Cephalic margin of apex of first gono- pod not constricted subdistally, termi- nating in acuminate projection (Fig. SB) sce. N. tencalanensis Campos, 1992 17. Laterocaudal side of apex of first gon- opod expanded (Fig. 5C)........... N. boyacensis Rodriguez, 1980 — Laterocaudal side of apex of first gon- opod not expanded 18. Outer laterocaudal surface of apex of . first gonopod with small blunt spines (Fig. 5D) N. lasallei Rodriguez, 1980 — Outer laterocaudal surface of apex of first gonopod with large, sharp spines (Fig. 5E) ... N. niceforoi (Schmitt, 1969) Acknowledgments This study was made possible, in part, by a visitor grant awarded to one of us (MRC) by the Smithsonian Institution’s Office of Fel- lowships & Grants. Illustrations were pre- pared by Juan C. Pinzén; Molly K. Ryan helped in the final preparation of the art work. Literature Cited Bott, R. 1967. Fliiss-krabben aus dem westlichen Siid- amerika.—Senckenbergiana Biologica 48(5/6): 365-372. 907 Campos, M. R. 1992. New species of fresh-water crabs of the genus Neostrengeria Pretzmann, 1965 (Crustacea: Decapoda: Pseudothelphusidae) from Colombia.—Proceedings of the Biological Society of Washington 105(3):540—554. . 1994. Diversidad en Colombia de los cangre- jos del género Neostrengeria.—Academia Col- ombiana de Ciencias Exactas, Fisicas y Natur- ales. Coleccio6n Jorge Alvarez Lleras 5:1—143. , & G. Rodriguez. 1985. A new species of Neostrengeria (Crustacea: Decapoda: Pseudoth- elphusidae) with notes on geographical distri- bution of the genus.—Proceedings of the Bio- logical Society of Washington 98(3):718—727. Milne Edwards, H. 1853. Mémoire sur la famille des Ocypodiens.—Annales des Sciences Naturelles, Zoologie 20:163—228. Pretzmann, G. 1965. Vorlaufiger Bericht tiber die Fam- ilie Pseudothelphusidae.—Anzeiger der Oster- reichischen Akademie der Wissenschaften Mathematische Naturwissenschaftliche Klasse (1), 1:1-10. Rathbun, M. 1893. Descriptions of new species of American freshwater crabs.—Proceedings of the United States National Museum 16(959): 649-661, pl. 73-77. . 1897. Descriptions de nouvelles espéces de crabes d’eau douce appartenant aux collections du Muséum d’ Histoire naturelle de Paris.—Bul- letin du Muséum National d’ Histoire Naturelle, Paris 3(2):58—61. Rodriguez, G. 1980. Description préliminaire de qu- elque espéces et genres nouveaux de crabes d’eau douce de |’Amérique tropicale (Crusta- cea, Decapoda, Pseudothelphusidae).—Bulletin du Muséum National d’ Histoire Naturelle, Paris 4(3):889—894. . 1982. Les crabes d’eau douce d’ Amérique. Famille des Pseudothelphusidae.—Faune Trop- icale 22:1—223. . 1985. A new cavernicolous crab (Crustacea, Decapoda, Pseudothelphusidae) from Colom- bia.—Bioloski vestnik, Ljubljana 33(2):73—80. , & M. Tiirkay, 1978. Der generische Status einiger Kolumbianischer Stisswasserkrabben.— Senckenbergiana Biologica 59:297—306. Schmitt, W. 1969. Colombian freshwater crab notes.— Proceedings of the Biological Society of Wash- ington 82:93—112. Smalley, A. 1964. A terminology for the gonopods of the American river crabs.—Systematic Zoology 13:28-31. Smithe, E B. 1975. Naturalist’s color guide. The Amer- ican Museum of Natural History, New York. Part 1:unnumbered pages. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):908—-911. 1998. A new species of mud shrimp, Upogebia toralae, from Veracruz, México (Decapoda: Thalassinidea: Upogebiidae) Austin B. Williams and Jorge L. Hernandez-Aguilera (ABW) National Marine Fisheries Service Systematics Laboratory, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (JLH-A) Facultad de Ciencias e Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de México, A.P. 70306, México, 04510. D.F Abstract.—Upogebia toralae, a new species of mud shrimp from Veracruz, México is described and illustrated. The unique female holotype was collected intertidally in the Port of Veracruz which is strongly impacted by human ac- tivity. The species shares with many members of the genus from the western hemisphere a strongly developed proximal mesioventral spine on the merus of the second pereopod. The species stands alone, however, with respect to other characters. The triangular rostrum has no ventral spines, the anterior gastric region bears many anteriorly setose spines that are more or less transversely elongate and scalelike, and the palms of the chelae have a dorsal ridge that bears three erect and somewhat hooked spines on its proximal end. Infaunal crustaceans collected from the State of Veracruz, México in the south- western Gulf of México are known to in- clude a wide variety of decapod crustacean species (Hernandez-Aguilera et al. 1996). This species richness has been attributed to the location of Veracruz in the tropical zone, with wet weather and abundant rains, where temperature varies from 10° to about 35°C (Soto & Garcia 1989). Nearshore hab- itats include a great variety of biotopes such as sandy beaches, estuaries, extensive man- grove swamps, marshes, and nearly 28 cor- al reefs. A mud shrimp, Upogebia toralae, new species, has been collected in intertidal waters of this environment in an area of the Port of Veracruz that is strongly impacted by human activity, including oil spills and industrial pollution, urban effluents, and tourism. Upogebia toralae, new species Fig. 1 Material examined.—México: USNM 285522, 1 2 (holotype), S of Puerto de Ve- racruz, Ver. México, 19°11'43", 96°07'36"W, intertidal, 8 May 1997, col. R. E. Toral-Al- mazan and J. L. Hernandez-Aguilera. Diagnosis.—Projections to either side of rostrum each ending in a spine; single spine on postocular margin; anterior gastric re- gion bearing many anteriorly setose spines more or less transversely elongate and scalelike. Abdominal sternites unarmed. Telson subrectangular. Merus of cheliped bearing subdistal dorsal spine and row of 5 spines on ventral margin; carpus with strong mesiodistal dorsal spine and 2 mod- erate spines on mesiodistal margin; palms of chelae bearing 3 erect and somewhat hooked spines on proximal end of dorsal ridge. Merus of pereopod 2 with proximal mesioventral spine and 1 subdistal dorsal spine; carpus with 1 subdistal dorsal spine and 1 distoventral spine. Merus of pereopod 3 with 1 distodorsal spine and row of 3 spines on ventral margin. Merus of pereo- pod 4 spineless. Description.—Rostrum triangular, hori- zontal in lateral view but with slightly downturned tip exceeding eyestalks by in- VOLUME 111, NUMBER 4 Fig. 1. 909 Upogebia toralae, new species, USNM 285522, 2 Holotype; a, carapace and cephalic region, lateral; b, anterior carapace, dorsal; c, telson and uropods, and part of abdominal segment 6, dorsal; d, cheliped, right lateral; e, cheliped, right mesial; f/ pereopod 2, left; g, pereopod 3, right; h, pereopod 4, left (fragment); i, pereopod 5, right. Scale = 3 mm. terval equal to length of cornea; 2 subdistal erect dorsal spines with tiny corneous tips followed on each side by smaller marginal spines, 3 on right, 4 on left; median line raised into low ridge. Pilose-armed field on anterior gastric region of carapace orna- mented with rather sparse, transversely elongated, well separated scalelike spines bearing short setae anteriorly, about 10 of these spines along each lateral margin of anterior gastric region; scattered smaller scalelike spines mesial to each lateral mar- gin, posterior region of carapace glabrous. Lateral ridge on either side of anterior gas- tric region extended anteriorly into process lateral to rostrum and bearing crest of 9 and 11 spines, more on left than on right. Shoul- der lateral to cervical groove bearing about 4—5 spines below intersection with thalas- sinidean line, dorsal 2 spines prominent and 910 acute, spines anteroventral to these much smaller or obsolescent; thalassinidean line continuing to posterior margin of carapace without interruption; postocular margin of carapace armed with acute spine at level of eyestalk. Abdominal sternites unarmed. Telson subrectangular, posterior margin shallowly biarcuate, smooth; transverse proximal ridge prominent, lateral ridge at each side obsolescent. Eyestalk stout, slightly elevated distally, reaching along basal % of rostrum; lower margin slightly convex; cornea narrower than diameter of stalk and directed ventro- laterally. ~ Antennular peduncle reaching to about *% length of terminal article of antennal pe- duncle, combined length of proximal 2 ar- ticles subequal to length of terminal article. Antennal peduncle with distal article and distal half of penultimate article extending beyond tip of rostrum; article 2 bearing strong, slender, subdistal ventral spine; moderate oval scale bearing small upturned distal spine. Maxilliped 3 bearing epipod. Epistomal projection rather broad in lat- eral view, bearing prominent apical spine. Chelipeds with ventral margin of ischium bearing 1 spine. Merus with row of 5 spines on ventral margin and a subdistal spine on dorsal margin. Carpus trigonal, with shal- low longitudinal lateral groove, strong Spine at anterior ventrolateral corner; dorsal crest margin bearing only 1 spine on basal 4 of its length, obscured by oblique tuft of setae; short spine on anterodorsal margin mesial to articulation with propodus; 2 moderate spines on anteromesial margin. Chela length about 2.7 times chela height; palm with dorsal ridge bearing 3 prominent slightly hooked spines at its proximal end, paralleled by sparsely setose mesiodorsal ridge ending in small distal spine; ciliated oblique ridge on lower lateral surface; both lateral and mesial surfaces of palm obscure- ly punctate. Fixed finger with extended slender tip and lobular tooth on occlusive PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON edge. Dactyl at least twice length of fixed finger, drawn to corneous tip and bearing 2 lobular teeth on proximal half of occlusive edge, distalmost tooth opposing tip of fixed finger; dorsal and lateral longitudinal ridges heavily setose. Pereopod 2 reaching about to distal 4 of cheliped palm; carpus with acute subdistal dorsal spine and nearly equal subdistal ven- tral spine; merus with slender subdistal- spine on dorsal margin and very strong proximal mesioventral spine. Pereopod 3 with carpus bearing 2 somewhat hooked dorsal spines on proximal % of length; mer- us bearing distodorsal spine and 3 strong spines on ventral margin. Pereopod 4 (frag- mentary) with spineless merus. Pereopod 5 of usual form, with cleaning brush on pro- podus. Uropods with acute spine on protopod above base of mesial ramus; lateral ramus with mesial rib bearing smaller spine prox- imally; both rami slightly exceeding telson, and with distal margins bearing rather uni- formly spaced row of granules. Measurements (in mm).—Anterior car- pace length 5.3, carapace length 7.5, length of chela including fixed finger 3.5, mid- length height of chela 1.3. Known range.—Confined to type locality. Remarks.—Upogebia toralae, new spe- cies, shares with many eastern Pacific and western Atlantic members of the genus a second pereopod on which the merus bears a strong proximal mesioventral spine (see keys to species in Williams 1986, 1993). The abdominal sternites and pleura bear no ventral spinules. The species stands alone, however, with respect to several characters. The triangular rostrum has no ventral spines. The anterior gastric region bears many anteriorly setose spines that are more or less transversely elongated and scalelike rather than being simple spines or spinelike tubercles. The palms of the chelipeds have a dorsal ridge that bears 3 erect and somewhat hooked spines on its proximal end. Moreover, the carpus has a mesiodorsal crest that bears 1 VOLUME 111, NUMBER 4 spine on the basal % of its length as well as the usual strong distal spine rather than a series of spines along all of its length. In the key to species of Upogebia in the West- erm Atlantic (Williams 1993), U. toralae lies closest to U. marina Coelho. Etymology.—The species is named in honor of Rosa Estela Toral-Almazan, Fa- cultad de Ciencias, UNAM, who has con- tributed to many collections from the Gulf of Mexico, in one of which the holotype was taken. Acknowledgments We thank Alfredo Ruiz-Nufio, Darryl L. Felder, and Michael Vecchione for critical reading of the manuscript, and Mollie Or- emland for preparing the illustrations. This study was supported in part under funding 911 to one of us (JLH-A) through the Comisi6n Nacional para el Conocimiento y Uso de la Biodiversidad (FB384/HO22/97). Literature Cited Hernandez-Aguilera, J. L., R. E. Toral-Almazan, & J. A. Ruiz-Nufio. 1996. Especies catalogados de crustaceos estomat6podos y decapodos para el Golfo de México, Rio Bravo, Tamps. a Progre- so, Yuc. Secretaria de Marina y Comision Na- cional para el Conocimiento y Uso de la Bio- diversidad, México, D. F, 132 pp. Soto, M., & E. Garcia. 1989. Atlas climatico del Es- tado de Veracruz. Instituto de Ecologia, México, 125 pp. Williams, A. B. 1986. Mud shrimps, Upogebia, from the eastern Pacific (Thalassinoidea: Upogebi- idae).—Memoirs of the San Diego Society of Natural History 14:1—60. . 1993. Mud shrimps, Upogebiidae, from the western Atlantic (Crustacea: Decapoda: Thal- assinidae).—Smithsonian Contributions to Zo- ology 544:1-77. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):912-915. 1998. Pinnotheres malaguena Garth, 1948, a new member of the genus Fabia Dana, 1851 (Crustacea: Brachyura: Pinnotheridae) Ernesto Campos and Raymond B. Manning (EC) Facultad de Ciencias, Universidad Autonoma de Baja California, Apartado Postal 2300, Ensenada, Baja California, 22800 México; (RBM) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560-0163, U.S.A. Abstract.—The original description and figures of the male holotype of the pinnotherid crab Pinnotheres malaguena Garth are emended. The morphology of this species excludes it from Pinnotheres Bosc, 1802 but supports its transfer to the genus Fabia Dana, 1851. Fabia malaguena can be distinguished from other members of Fabia by its unique gonopod shape and its abdomen, which has somites 5 to the telson fused. The pinnotherid crab species Pinnotheres malaguena Garth, 1948 was described on the basis of an adult male collected in Ma- laga Bay, Colombia [Malaga = 6°44’N, 72°45'W] (Garth 1948). Gore (1986) and Campos (1996), based on Garth’s account, suggested that P. malaguena may belongs in the genus Fabia Dana, 1851. Our study of the male holotype, deposited in the American Museum of Natural History (AMNH), allows us to confirm this. The morphology of P. malaguena does not agree with Pinnotheres sensu stricto (see Manning 1993 for the characteristics of that genus), but it largely concurs with those of the genus Fabia. In addition to transferring P. malaguena to Fabia, we emend its original description (underlined in the text) and figures, since several mistakes and omissions in the orig- inal account were detected. Other abbrevi- ations used are WL = walking legs; MXP3 = third maxilliped. Fabia malaguena (Garth, 1948), new combination (Fig. 1) Pinnotheres malaguena Garth, 1948:53—S55, ine Se Pinnotheres malaguena.—Silas & Alagar- swami, 1967:1202, 1218.—Schmitt et al., 1973:56.—Gore, 1986:147.—Lemaitre & Alvarez Leon, 1992:61.—Hendrickx, 1995:142.—Campos, 1996:1161. Distribution.—Known only from the type locality, Malaga Bay, Colombia, in 4— 9 m depth (Garth 1948). Material examined.—Male holotype (AMNH 10012). Measurements.—Carapace length 2.6 mm, width 2.3 mm; frontorbital width 1.1 mm; length of chela 1.1 mm, of dactyl 0.5 mm; length of manus 0.6 mm (after Garth 1948). Redescription.—Carapace (Fig. 1A) slightly longer than broad, widest at middle, suboctagonal, angles rounded, dorsal surface smooth and bare, porcelain-like, strongly convex, without indications of regions, fron- tal and anterolateral margins clothed with shaggy hair-like setae. Front advanced con- siderably beyond orbits, its edges represent- ing a continuation of anterolateral margins interrupted only by sinuous indentation of orbits. Anterior margin subtruncate, appear- ing bilobed, edges concealed by fringing se- tae. Anterolateral margin longer than pos- terolateral, sloping at fairly steep angle from VOLUME 111, NUMBER 4 913 Fig. 1. Fabia malaguena (Garth, 1948), male holotype (AMNH 10012). Carapace length 2.6 mm, width 2.3 mm. A, Dorsal view; B, MXP3; C, Right chela, outer face; D, Abdomen; E, Gonopod. Setae omitted in Fig. 1B, D. the orbits, their gentle arching accentuated by a thick fringe of fur-like pile, longest me- dially. A suggestion of short, transverse line of hairs at gastric level. Posterior margin al- most straight, rimmed and bare. Orbits small and circular, eyestalks short, eyes filling sockets, corneas when retracted concealed by setae in dorsal view. Antenna short, basal article hiatus, flagellum only extending be- yond margin of front. 914 MXP3 (Fig. 1B) nearly transverse in po- sition, gently convex distally, carpus longer than wide, cylindrical, curving inward and broadening distally; propodus subequal to carpus in length, flattened and broadened medially, outer surface lacking neither row of setae nor transverse ridge as described and figured in original account; dactylus small, digitiform, placed in angular notch in middle third on ventral margin of propodus, and falling short of end of propodus. Chelipeds (Fig. 1C) stout, equal, merus and carpus fringed with setae above and be- low, leaving a smooth, bare space between; chela with similar open area, fringing setae as dense as those of anterolateral margins of carapace, forming superior crest. Fingers Slender, thin, tapering distally, sharply pointed, dactylus curving strongly down- ward, pollex little deflexed, both fingers with subtruncated tooth proximally placed on cutting edge, a few long setae visible in narrow gape, tips crossing until pointed in almost opposite directions. Sternal plastron flattened at center, slight- ly concave towards margins, latter fringed with setae, segmentation clearly indicated. Male abdomen (Fig. 1D) widest at third so- mite, sides of latter broadly rounded, taper- ing to narrowest point at middle of fused fifth somite to telson, then broadening slightly before the semicircular tip which is edged with fine setae (omitted in Fig. 1D). Gonopod (Fig. 1E) cylindrical, sickle shaped, tapering gradually from base al- most to gutter-like tip which flares slightly and divided longitudinally for short dis- tance along its length; margins setose, long plumose setae extending to base of append- age externally. WL (Fig. 1E) symmetrical, laterally com- pressed, with exception of dactyli, margins covered with fringe of furry setae; WLI twisted, WL 2-3 with two fringes of long swimming setae, one on outer face of car- pus and propodus, one on ventral margin of propodus. Meri subequal in shape and width; carpi trigonal, slender proximally, broadening distally; propodi widest at mid- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON point, obliquely rounded distally; dactyli long, tapering abruptly to sharp, transpar- ent, curved nails, prehensile edge smooth. Relative length of WL 2 >3 > 1 > 4. Remarks.—The following features sup- port the inclusion of the male of Pinno- theres malaguena in the genus Fabia: the carapace is convex, smooth, shiny, porce- lain-like; the frontorbital and anterolateral margins with stout, hair-like setae; MXP3 with a subtrapezoidal propodus subequal to the cylindrical carpus, and digitiform dac- tylus inserted in the middle third on the ventral margin of the former article; WL1 are twisted and margins of the compressed WLI1-—4 are covered with a fringe of hair- like setae; and the abdomen with somites 1—4 free, 5 to the telson fused. Males in the genus Pinnotheres have the dactylus of MXP3 styliform, inserting basally on the ventral margin of the propodus and all of the abdominal somites and telson free. Fabia malaguena can be separated from other species of the genus Fabia by its sin- gular gonopod shape (Fig. 1E) and because abdominal somites 5 to the telson are fused (Gigs iD): Acknowledgments We are indebted to Lara L. Tolchin, American Museum of Natural History, for the loan of the holotype of Pinnotheres ma- laguena (Garth, 1948); and to Alma Rosa de Campos for her fine artistic work. This work was partially supported by the project UABC-CONACyT 3587-N9311, and the program 0134-UABC “Systematics of the symbiotic crustaceans of the Mexican Pa- cific.”” This is contribution #447 from the Smithsonian Marine Station at Fort Pierce; support of that program for Manning’s stud- ies on pinnotherids is gratefully acknowl- edged. Literature Cited Bosc, L. A. G. 1802. Histoire naturelle des Crustacés, contenant leur description et leurs moeurs, avec figures dessinées d’aprés nature. Paris, 1:1—258, pls. 1-8; 2:1—296, pls. 9-18. VOLUME 111, NUMBER 4 Campos, E. 1996. Partial revision of the genus Fabia Dana, 1851 (Crustacea: Brachyura: Pinnotheri- dae).—Journal of Natural History 30:1157— 1178. Dana, J. D. 1851. On the classification of the Crustacea Grapsoidea.—American Journal of Science and Arts, series 2, 12:283—291. Garth, J. S. 1948. The Brachyura of the “Askoy”’ ex- pedition with remarks on carcinological col- lecting in the Panama Bight.—Bulletin of the American Museum of Natural History 92(1):1— 66. Gore, R. H. 1986. Fabia felderi species novum, a new pinnotherid crab from the central eastern coast of Florida (Crustacea: Decapoda: Brachyura).— Northeast Gulf Science 8:143-148. Hendrickx, M. E. 1995. Checklist of brachyuran crabs (Crustacea: Decapoda) from the eastern tropical Pacific.—Bulletin de 1’Institut Royal des Sci- ences naturelles de Belgique, Biologie, 65:125— 150. Lemaitre, R., & R. Alvarez Leon. 1992. Crustdceos 915 decapodos del Pacifico Colombiano: lista de es- pecies y consideraciones zoogeograficas.—An- ales del Instituto de Investigaciones Marinas de Punta de Betin 21:33-76. Manning, R. B. 1993. West African pinnotherid crabs, subfamily Pinnotherinae (Crustacea, Decapoda, Brachyura.—Bulletin du Muséum national d’Histoire naturelle, series 4, 15(A, 1—4):125— 177. Silas, E. G., & K. Alagarswami. 1967. On an instance of parasitisation by the pea-crab (Pinnotheres sp.) on the backwater clam [Meretrix casta (Chemnitz)] from India, with a review of the work on the systematics, ecology, biology and ethology of pea crabs of the genus Pinnotheres Latreille.—Proceedings of the Symposium on Crustacea held at Ernakulam from January 12 to 15, 1965. Marine Biological Association of India, Symposium series 2, 3:1161—1227. Schmitt, W. L., J. C. McCain, & E. S. Davidson. 1973. Family Pinnotheridae. Decapoda I, Brachyura I. In H. E. Gruner & L. B. Holthuis, eds. Crusta- ceorum Catalogus 3:1—160. Den Haag, W. Junk. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):916—920. 1998. Cave chaetognaths in the Canary Islands (Atlantic Ocean) F Hernandez and S. Jiménez Museo de Ciencias Naturales (Dpto. de Biologia Marina), Apto. 853. 38080, Santa Cruz de Tenerife, Islas Canarias Abstract.—Morphological and biometric observations of chaetognaths found in a submarine cave on the southeast coast of Tenerife (Canary Islands) are presented. The specimens, that we have called Spadella aff. ledoyeri, differ notably from the coastal species Spadella cephaloptera, which is a well known inhabitant of the submerged seagrass beds of the infralittoral areas of the is- lands. Because the genus Paraspadella was previously described from an an- chialine cave on the Grand Bahama Island (western Atlantic Ocean), this is the second record of an Atlantic cave chaetognath and the first for the genus Spa- della in Atlantic Ocean caves. The Canary Island benthic chaetognaths, unlike the pelagic species, have only rarely been studied. Among the few studies done are those of Hernandez & Jiménez (1992) and Broerse (1993), which deal with bio- metric aspects of Spadella cephaloptera (Busch, 1851) in the island of Tenerife. This species has been found in beds of the submerged sea grass Cymodocea nodosa and on patches of the alga Caulerpa proli- fera in shallow infralittoral areas of the is- lands of Tenerife (Hernandez & Jiménez 1992, Broerse 1993) and Gran Canaria, es- pecially in areas with a sandy substratum. The discovery of benthic cave chaetognaths in the Mediterranean (Casanova 1986, 1992) and in the Bahamas (Bowman & Bieri 1989) has led us to trawl for them in dark submarine caves. Specimens of these organisms were found, but they were dif- ferent from descriptions of the benthic spe- cies S. cephaloptera, which is well known in the islands. Materials and Methods The material collected comes from a dark, submarine cave, situated on the south- east coast of the island of Tenerife (Canary Islands) (Fig. 1), between the towns of San- ta Cruz and Candelaria. Two trawls were carried out, using a 200 ym, manual plank- ton net, 12 m from the entrance and a few cm above the bottom. A total of 42 chae- tognath specimens belonging to the genus Spadella was obtained. The specimens were measured to give data on total length, caudal length, ovarian length, and the number of grasping spines and teeth. For comparison, we used speci- mens of Spadella cephaloptera, which were caught on the same dates as the cave spe- cies. These were caught in trawls over beds of sea grass and algae off the islands of Tenerife and Gran Canaria. Cave Characteristics The cave is a volcanic pipe at the bottom of a submarine canyon. It has a broad en- trance and a sandy floor. The cave narrows close to the entrance and then widens out again after about one m. The back of the cave is marked by a natural blockage of sand. The cave is 16 m below ground level, with a slope of 1.3 m and is 15 m deep. Geological interest in the cave stems from the fact that it forms part of the lava fields that flowed from the back of La Esperanza (Tenerife) to the coast and into the sea. VOLUME 111, NUMBER 4 - }- 29°N La Palma \} Tenerife La Gomera ~~ j- 28° El Hierro 18° 17° 16° | | | Canary Islands Gran Canaria 917 Lanzarote | ) ce} Fuerteventura 15° 14°0 [| | | Fig. 1. Apart from the chaetognaths, other fauna found in the cave included shrimps (spawn- ing females of Plesionika narval), tubicular Polychaeta, sponges, formations of Nadra- cis asperula (corals), prawns (Lysmata grabhami) and echiurids (Bonellia viridis). Results and Discussion The chaetognaths studied showed the fol- lowing taxonomic characteristics (Figs. 2, 3): General aspect of the body: white, long, thinner than in Spadella cephaloptera, which is thicker and yellowish brown. Head Square, not oval as in S. cephaloptera (Fig. 2). Short lateral fins (only 50% of the cau- dal region), wider than in S. cephaloptera, which start just before the tail. Ocular pig- ment present. The ciliata corona is com- pletely circular and different from that of S. cephaloptera studied in Tenerife and Gran Canaria, where it is more flattened and bi- lobed (Fig. 3). Caudal fin completely tri- angular; in S. cephaloptera, it is spatula shaped. Triangular seminal vesicles, sepa- rated from the lateral and caudal fins. In S. cephaloptera, the seminal vesicles touch both fins. Caudal region strong and broad. Cephalic tentacles and intestinal diverticu- lae were not observed. Ovaries with large, rounded ovules. Ten or eleven reddish-light brown grasping spines. Only 3—4 anterior Location of the sample station in the southeast of Tenerife (Canary Islands). teeth observed. The maximum size for stage III sexual maturity is 4.5 mm (4.3 mm, mean) (Table 1). Biometric data is also provided for each sexual stage of the spec- imens of this study (Table 1). These data have been compared with the specimens corresponding to the Tenerife (Hernandez & Jiménez 1992) and Gran Canaria speci- mens (Table 2). The specimens collected in the cave be- long to the genus Spadella, but do not pre- sent the typical characteristics of Spadella cephaloptera. The specimens studied are closer to Spadella ledoyeri, a cave dwelling chaetognath of the Mediterranean (Casa- nova 1986). Among the morphological dif- ferences between our specimens and those of S. cephaloptera, we observed: the shape of the ciliata corona, shape and position of seminal vesicles, shape, position and exten- sion of the lateral fins, shape and general aspect of the body, shape and size of the collarette and the aspect of the eye pigment. Although the shape of the ciliata corona can vary (Ghirardelli 1968), the characteristics are valid and have a high taxonomic value. Concerning Atlantic cave-dwelling chae- tognaths, the only other species on record is Paraspadella anops, described from a single specimen from Sagittarius Cave near Gran Bahama Island (Bowman & Bieri 918 2 mm Fig. 2. 1989). This species, as in the case of our specimens of Spadella ledoyeri, is clearly the descendent of Paraspadella schizop- tera, the common neritic species of the re- gion, from which it differs in only a few characteristics. Likewise, Spadella ledoyeri is believed to have evolved from either Spadella ce- phaloptera, isolated in the caves of the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2.4 mm Selected specimens from the samples. A, Spadella aff. ledoyeri; B, Spadella cephaloptera. Mediterranean from submergence during the most recent ice age, or alternatively, from the deep water species, Spadella bi- rostrata, isolated in cavities of deeper caves flooded during a prior glaciation (Casanova ISS). For the first time in the Atlantic Ocean, benthic chaetognaths of the genus Spadella from underwater caves are described. In the Table 1.—Biometric characteristics of stage III specimens of Spadella aff. ledoyeri on Tenerife. TL = total length, CL = caudal length, %C/T = relative tail length in relation to total length. TL (mm) Stage I 3.0 n= 1 Stage II 3.04.5 n = 23 average: 3.9 Stage I 4.0-4.5 n= 18 average: 4.3 CL (mm) %oC/T 1.5 50.0 1.5—2.5 42.9-55.6 average: 2.0 average: 50.0 2.0-2.5 44.4-55.6 average: 2.2 average: 51.5 VOLUME 111, NUMBER 4 919 — Fig. 3. Spadella aff. ledoyeri. A, caudal fin; B, seminal vesicle; C, ocular pigment; D, head and neck. Spadella cephaloptera. E, caudal fin; K seminal vesicle; G, head and neck. 920 Table 2.—Comparison of total length (TL) for stage III specimens of Spadella aff. ledoyeri from Gran Ca- naria and Tenerife. TL (mm) Island Month Habitat 5.5 Gran Canaria April seagrass 4.0 Tenerife April seagrass 3.5. Tenerife December seagrass 4.5 Tenerife April cave Canary Islands, therefore, as is the case in the Mediterranean (Casanova 1992, genus Spadella) and the Bahamas (Bowman & Bieri 1989, genus Paraspadella), these an- imals exist in the biotope of submarine caves, where, according to the above men- tioned authors, there are original popula- tions. New collections will be done, at different times of the year, extending the study to other caves in the islands, in order to in- vestigate the biological cycle and to clarify certain aspects, which are debated in the Mediterranean, of the origin of the cave dwelling species and make accurate com- parisons between Spadella aff. ledoyeri and the coastal species Spadella cephaloptera from waters surrounding the Islands. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Acknowledgments Our sincere thanks go to Dr. Stephen Gardiner (Associate Editor) and to the col- lector, Alfredo Lainez, who sent us the specimens, together with collection data. Literature Cited Bowman, T. E., & R. Bieri. 1989. Paraspadella anops, new species, from Sagittarius cave Gran Ba- hama island, the second troglobitic chaeto- gnath.—Proceedings of the Biological Society of Washington 102:586—589. Broerse, A. T. C. 1993. Biometric variation in Spadella cephaloptera on the Canary islands (Chaeto- gnatha).—Beaufortia 43(6):101—113. Casanova, J. P. 1986. Spadella ledoyeri, chaetognathe nouveau de la grotte sous-marine obscure des Trémies (Calanque de Cassis).—Rapports Com- mision international Exploration Mer Méditer- ranée 30,2:196. . 1992. Les chaetognaths cavernicoles de la Méditerranée nord-occidentale: adaptations et spéciation, comparaison avec | Atlantique.— Bulletin Institute Océanographique Monaco 9: 83-100. Ghirardelli, E. 1968. Some aspects of the biology of the Chaetognaths.—Advances in Marine Biol- ogy 6:271—375. Hernandez, E, & S. Jiménez. 1992. Primeras obser- vaciones sobre la presencia del género bentdn- ico Spadella (Chaetognatha) en la isla de Ten- erife (Canarias).—Actas del V Simposio Ibérico de Estudios del Bentos Marino 2:95-102. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):921—928. 1998. The discovery of Glyphocrangon stenolepis Chace (Decapoda: Caridea: Glyphocrangonidae) from Taiwan and Japan, with notes on individual variation Tomoyuki Komai, Tin-Yam Chan, and Ding-An Lee (TK) Department of Zoology, Natural History Museum and Institute, Chiba, 955-2 Aoba-cho, Chuo-ku, Chiba 260, Japan; (TYC) Institute of Marine Biology, National Taiwan Ocean University, Keelung, Taiwan, R.O.C.; (DAL) Taiwan Fisheries Research Institute, 199 Ho-Ih Road, Keelung, Taiwan, R.O.C. Abstract.—Abundant material of Glyphocrangon stenolepis Chace was found off Taiwan. Supplementary specimens from the South China Sea and Japan have made possible to redescribe this poorly known species. Remarkable sexual differences in body sculpture and coloration are found in this species. The paratype from the Sulu Sea has proven to represent a different as yet undetermined species. Other morphological variations are discussed, and the coloration is illustrated. In his report on the glyphocrangonid and crangonid shrimps collected by the Philip- pine Albatross Expedition 1907—1910, Chace (1984) described Glyphocrangon stenole- pis, based on two specimens, one from the South China Sea, off Pratas Islands, and an- other from north of the Sulu Sea, Philip- pines. The specimen from the South China Sea is the holotype. The paratype from the Sulu Sea is much smaller than the holotype, and appears to be a juvenile or a young fe- male. Although Chace (1984) noted some differences between the holotype and par- atype, he interpreted them as size-related. The species has not been reported since the original description. Recent investigations by one of us (TYC) has shown that Glyphocrangon stenolepis is common in offshore waters of Taiwan. Fur- thermore, scientists from the Fisheries Re- search Institute of Taiwan collected four specimens from the type locality, off Pratas Islands in 1996. Through the kind courtesy of Prof. K. Hayashi, a single specimen from the East China Sea, off Japan, has also been made available for examination. All this material has enabled us to diagnose the spe- cies more precisely, and to extend its known geographical range northwards to Japan. The study of the material has shown that the species exhibits marked variation in body sculpture in males. Therefore, we pro- vide a full redescription of the species. Some important differences between the holotype and paratype noted by Chace (1984) indicate that the paratype represents another species, though its exact identity is still uncertain. The specimens, all collected by bottom trawls, are deposited in institutions indicat- ed by the following abbreviations: CBM, Natural History Museum and Institute, Chi- ba; NUE National University of Fisheries, Shimonoseki; NTOU, National Taiwan Ocean University, Keelung; TFRI, Taiwan Fisheries Research Institute, Keelung Branch; USNM, National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C. The terminology for the carinae and spines on the carapace follows Holthuis (1971) and Chace (1984). The ab- breviation cl indicates postorbital carapace length. 922 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. (net 0) Nn) yysr-nAAUK i @ ™))) Glyphocrangon stenolepis Chace, 1984. Animals in dorsal view, pereopods omitted. A, male from off Ta-Shi, NE Taiwan, cl 14.1 mm, CBM-ZC 3614; B, female from off Su-Aou, NE Taiwan, cl 14.7 mm, CBM-ZC 2935. Scale bar indicates 5 mm. Glyphocrangon stenolepis Chace, 1984 (Figs. 1—4) Glyphocrangon stenolepis Chace, 1984:22 (part), fig. 5 (not fig. 6; =Glyphocrangon Sp.). Type material.—South China Sea. Pratas Islands (Tungsha Tao), Albatross Stn 5300, 20°31’'N, 115°49’E, 485 m, 8 Aug 1908, 1 male (cl 11.3 mm), holotype (USNM 205091). Other material.—South China Sea. R.V. Fisheries Research I, 19°49.2'N, 114°09.3’E, 512 m, 23 Apr 1996, 1 male (cl 11.1 mm) (TFRI); exact position unknown, 1996, 3 males (cl 13.6—-14.8 mm) (TFRI). Taiwan. Fishing pots, commercial trawl- ers, 300-500 m, sandy mud bottoms: Ta- Fig. 2. A, male from off Ta-Shi, NE Taiwan, cl 14.1 mm, CBM-ZC 3614; B, female from off Su-Aou, NE Taiwan, cl 14.7 mm, CBM-ZC 2935. Scale bar indicates 5 mm. Glyphocrangon stenolepis Chace, 1984. Animals in lateral view, pereopods and pleopods omitted. VOLUME 111, NUMBER 4 923 BCDFHK G 2mm ene bonnes EGIJLN = 0.5mm Fig. 3. Glyphocrangon stenolepis Chace, 1984. Male from off Ta-Shi, NE Taiwan, cl 14.1 mm, CBM-ZC 3614. Left appendages. A, antenna, ventral, setae omitted; B, third maxilliped, lateral; C, first pereopod, lateral; D, second pereopod, lateral; E, chela of same, lateral; E third pereopod, lateral; G, tip of dactyl of same, flexor; H, fourth pereopod, lateral; I, tip of dactyl of same, flexor; J, same, lateral; K, fifth pereopod, lateral; L, tip of dactyl of same, flexor; M, endopod of first pleopod, ventral; N, appendices interna and masculina of second pleopod, mesial. 924 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Glyphocrangon stenolepis Chace, 1984. A, ovigerous female from Taiwan; B, male from Taiwan; C, male from Pratas Islands, South China Sea. VOLUME 111, NUMBER 4 Shi, I-Lan County (northeastern coast), 25 Nov 1994, 3 males (cl 12.1—13.2 mm), 4 ovig. females (cl 12.2—-14.5 mm) (NTOU); 27 Apr 1995, 1 ovig. female (cl 13.9 mm), 1 female (cl 15.4 mm) (NTOU); 13 June 1995, 2 males (cl 13.3, 14.5 mm), 1 ovig. female (cl 15.0 mm) (CBM-ZC 3614); 3 July 1995, 2 ovig. females (cl 13.6, 14.6 mm); 11 Mar 1997, 18 males (cl 11.4—16.1 mm), 35 ovig. females (cl 12.3—-16.0 mm), 12 females (cl 11.0—16.0 mm) (NTOU); 10 males (cl 11.8—16.7 mm), 7 ovig. females (cl 13.6—-17.5 mm), 3 females (CL 13.0— 13.4 mm) (CBM-ZC 3918); 25 Feb 1997, 1 ovig. female (cl 14.2 mm) (NTOU); 4 Dec 1997, 1 female (cl 11.7 mm) (CBM- ZC 3882); 1 June 1998, 3 males (cl 9.2— 13.0 mm), 1 ovig. female (cl 15.7 mm) (NTOU). Su-Aou, I-Lan County (northeast- ern coast), 20 Apr 1985, 5 males (cl 12.4— 15.0 mm), 4 ovig. females (cl 13.0—16.0 mm) (CBM-ZC 3615); 2 May 1985, 4 males (cl 12.5—13.1 mm), 5 ovig. females (cl 12.4-13.3 mm), | female (cl 11.4 mm) (NTOU); 6 Aug 1996, 1 male (cl 13.9 mm), 2 females (cl 14.7, 15.3 mm) (CBM-ZC 2935); 7 Aug 1996, 1 male (cl 16.6 mm) (NTOU); 18 Nov 1997, 1 male (cl 9.5 mm) (NTOU); 5 Dec 1997, 7 males (cl 10.3— 14.4 mm), 1 ovig. female (cl 15.2 mm) (CBM-ZC 3901). Tong-Kong, Ping-Tong County (southwestern coast), 25 Feb 1995, 1 male (cl 12.5 mm) (NTOU); 30 May 1997, 1 male (cl 10.5 mm) (NTOU). East China Sea. R. V. Tennyo-Maru, west of Tokara Islands, southern Japan, Stn T-2, 29°20'N, 127°26’E, 488 m, 1 male (cl 12.8 mm) (NUF). Redescription.—Integument not pubes- cent. Rostrum (Figs. 1, 2) curved dorsad anteriorly, 0.95—1.23 times as long as car- apace, armed with 2 pairs of lateral teeth, posterior pair (arising from posterior to lev- el of posterior margin of orbit) more or less reduced, sometimes barely discernible, nev- er acute; series of distinct transverse septa on anterior part of dorsal surface, median carina present on distal portion and poste- rior portion between eyes. Carapace (Figs. 925 1, 2) with anterior first (Submedian) carina varying from slightly uneven to distinctly 5-lobed; median area anterior to submedian carina with single tubercle in midline; pos- terior first (submedian carina) composed of 2 elongate ridges. Anterior second (inter- mediate) carina composed of 3 obtuse lobes; posterior second (intermediate) cari- na also 3- or 4-lobed. Intercarinal space be- tween posterior first and second carinae usually with row of tubercles. Hepatic re- gion with 2 or 3 tubercles; posterior third (antennal) carina 3- or 4-lobed. Anterior fourth (lateral) carina forming bilobate, wing-like expansion, terminating anteriorly in sharp point independent of branchioste- gal spine, arising from posterior to level of orbit, posterior lobe in line with non-dentate anterior portion; posterior fourth (lateral) carina interrupted posteriorly, most part nei- ther dentate nor lobate. Anterior fifth (sub- lateral) carina not prominent, nearly linear; posterior fifth (sublateral) carina very short. Sixth (submarginal) carina absent anterior- ly, indistinct posteriorly. Margins of carinae faintly erose. Antennal spines unarmed marginally, less than 0.5 as long as and di- verging littke more than anteriorly directed branchiostegal spines. Abdomen (Figs. 1, 2) with ridges and tu- bercles on dorsal surface blunt or obsolete; prominence of ridges and tubercles variable in males, well developed in females. First somite with distinct submedian tubercles on posterior section of tergum in females, sometimes faint in males; median carina obsolete to distinct; transverse groove mod- erately shallow to faint. Broad median, sub- median and lateral carinae on second and third somites interrupted by transverse groove, distinct in females, sometimes ob- solete or absent in males; median carina on fourth somite distinct in both sexes, more or less notched at anterior 0.3; fifth somite with prominent anterior tubercle and sharp carina posteriorly along midline, latter flanked by posteriorly divergent sharp ca- rinae; sixth somite with median carina di- vided in 2 by shallow, sometimes very 926 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Glyphocrangon stenolepis Chace, 1984. Branchial formula. Maxillipeds 1 2 3 Pleurobranchs = _— es Arthrobranchs — — Podobranch — = ae Epipods 1 1 = Exopods 1 weak, notch anteriorly. Second somite with pleuron bearing 2 short, sometimes blunt teeth and inconspicuous anterior lobe; pleu- ra of third, fourth and fifth somites each with 2 moderately short marginal teeth. Eyes (Figs. 1, 2) moderately large for ge- nus, cornea lightly pigmented. Antennular peduncle (Figs. 1, 2) over- reaching distal margin of scaphocerite by 0.2—-0.3 length of intermediate segment; outer flagellum longer and thicker in males than in females. Scaphocerite (Fig. 3A) elongate oval, 1.95—2.37 times longer than wide, with small lateral tooth arising slight- ly posterior to level of mid-length, marginal setae confined to blade distal to lateral tooth; carpocerite not overreaching distal margin of blade. Mouthparts typical of species in genus. Third maxilliped (Fig. 3B) moderately stout, reaching or overreaching distal mar- gin of scaphocerite; antepenultimate seg- ment distinctly carinate dorsolaterally. First pereopod (Fig. 3C) incompletely subchelate; ischium with distoventral por- tion strongly produced. Second pereopods (Fig. 3D) nearly equal, each with 23-27 carpal articles; chela (Fig. 3E) flattened, with short fixed finger and strongly oblique cutting edge. Third to fifth pereopods (Fig. 3E H, K) moderately slender, each with subspatulate dactyl and merus longer than carpus and propodus combined. Third pe- reopod (Fig. 3F) with dactyl 0.4—0.5 times as long as propodus, terminating in simple unguis (Fig. 3G); propodus with terminal or subterminal setae. Fourth pereopod (Fig. 3H) with dactyl 0.6 times as long as pro- podus, terminating in small lobe mesiad to Pereopods small subterminal spine, bearing few short bristles on extensor surface distally (Fig. 3J); propodus with distal setae. Fifth pereo- pod (Fig. 3K, L) generally similar to fourth pereopod; dactyl 0.4—0.5 times as long as propodi, lacking bristles on extensor sur- face. Thoracic sternite deeply depressed in both sexes, anterior part of sixth thoracic sternite produced anteriorly as subtriangular lobe; eighth sternite posteriorly with obtuse median tubercle in males, unarmed in fe- males. Interlocking mechanism of carapace and thoracic sternum well developed. First pleopod of male with well-devel- oped appendix interna on endopod (Fig. 3M), bearing rows of long setae basally. Appendix masculina of second pleopod (Fig. 3N) reaching or slightly overreaching appendix interna, bearing numerous long bristles terminally and mesially. Branchial formula as shown in Table 1. Eggs large and elongate oval, ranging from 1.2 to 1.5 and 2.2 to 2.5 mm in short axis and long axis respectively. Coloration.—Females with body orange brown, ridges and carinae somewhat orange to reddish. Antennular and antennal flagel- la, distal parts of pereopods, posterior mar- gin of abdominal tergites and posterior parts of tail-fan reddish. Meri of posterior pereopods and ventral parts of abdominal pleura slightly whitish. Eyes golden brown. Eggs blue, and developed ovaries visible in- side carapace deep blue. Males with body light brown to slightly whitish, and gener- ally with color paler than females, particu- VOLUME 111, NUMBER 4 larly those with less developed abdominal sculpture. Size.—Males: cl 9.2—-16.7 mm; females: cl 11.0—17.5 mm; ovigerous females: cl 12.2—17.5 mm. Distribution.—East and South China Seas; 300-512 m; inhabiting soft bottoms. Remarks.—As previously mentioned, Glyphocrangon stenolepis was described based on two specimens, the holotype from off Pratas Islands, South China Sea, and the paratype from the Sulu Sea, the Philippines. The present material extends its geograph- ical range to southern Japan. Furthermore, this species is a common by-catch of the deep-sea commercial trawlers in Taiwan. This study has shown that the sculpture of the carapace and abdomen of the species is generally similar in females, whereas it is rather variable in males. In females, teeth, tubercles, carinae and grooves on the cara- pace and body are distinct (Figs. 1B, 2B), whereas in males, these structures are fre- quently obsolete to faint (Figs. 1A, 2A), or occasionally as distinct as in females. Also, there seems to be a relationship between body color and degree of development of abdominal sculpture in this species. The coloration of females is always orange to orange brown (Fig. 4A). In males, however, those with distinct abdominal sculpture have a similar coloration as in females, al- though those with eroded sculpture have a much paler coloration (Fig. 4B), and those of intermediate sculpture have intermediate coloration. The holotype represents an ex- ample of variation with least developed body sculpture, with short longitudinal ridges on the anterior abdominal tergites completely eroded, and the median carina of the fourth abdominal tergite not inter- rupted (see Chace 1984:fig. 5). Although the least sculptured Taiwanese specimens still have some traces of short longitudinal ridges on the abdominal tergites (Fig. 1A) and a small notch on the median carina of the fourth abdominal tergite, the abdominal sculpture of the three larger specimens from the type locality Pratas Islands (similar to 927 those shown in Figs. 1A, 2A) fit well within the range observed in the Taiwanese mate- rial. In addition, the coloration of the small Pratas male is very similar to that of some males from Taiwan (Fig. 4B, C). Therefore, it is concluded that the Taiwanese and South China Sea specimens are conspecific. Nevertheless, it seems that for similar size males, those from Taiwan are generally more sculptured than those from the South China Sea. Perhaps more specimens, partic- ularly females, from the South China Sea, will provide better insights on the geo- graphical variations of this species. In ad- dition to the variability of the body sculp- ture, the material reported herein displays variation in other important features: the proportional length of the rostrum varies from 0.95 to 1.23 times as long as the car- apace; the posterior pair of the lateral ros- tral teeth are sometimes barely discernible, as noted by Chace (1984), or are sometimes more prominent, showing as dentiform tu- bercles (Fig. 2B); the posterior fourth carina is usually interrupted at near the posterior end, and is rarely continuous. As mentioned by Chace (1984), the elon- gate oval antennal scaphocerite, the config- uration of the anterior fourth (lateral) cari- na, the long branchiostegal spine and the septate rostrum immediately separate G. stenolepis from most other species of the genus. Chace (1984) also noted that the ju- venile or female paratype was different from the holotype in the more outstanding anterior tooth on the anterior fourth (lateral) carina, the more strongly lobate posterior first, second and third carinae, and the more strongly sculptured abdomen. Except for the sculpture of the abdomen, the present study found that those differences are con- stant between the paratype and the present material of G. stenolepis, not associated with growth as Chace suggested. The small- est specimen in the present material is a male of cl 9.2 mm from Taiwan, slightly larger than the paratype (cl 8.1 mm, USNM 205091). The smallest Taiwanese male has a well developed abdominal sculpture as in 928 females, and the appendix masculina of the second pleopod is only slightly shorter than the appendix interna. Therefore, if the par- atype belongs to the same species as the holotype, it should be a female. However, none of the abundant females examined in the present study show a prominent anterior tooth on the anterior fourth carina as in the paratype. In addition to above differences, the posterior part of the third abdominal so- mite is more strongly elevated in the para- type than in the holotype or all other spec- imens of G. stenolepis. These differences indicate that the paratype represents a dif- ferent species. More material from the Sulu Sea is needed to determine the exact iden- tity of Chace’s (1984) paratype. Acknowledgments We are most grateful to E A. Chace, Jr, and R. B. Manning, National Museum of PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Natural History, Smithsonian Institution, Washington, D.C., for sending us the type material of Glyphocrangon stenolepis on loan, and Prof. K. Hayashi of National Fishery University, Shimonoseki, for mak- ing available the specimen of the species from Japanese waters for study. The present study is supported by a research grant on the decapod crustaceans of Taiwan from the National Science Council, Taiwan, R.O.C. Literature Cited Chace, E A., Jr., 1984. The caridean shrimps (Crusta- cea: Decapoda) of the Albatross Philippine Ex- pedition, 1907—1910, Part 2: Families Glypho- crangonidae and Crangonidae.—Smithsonian Contributions to Zoology 397:i-1v, 1—63. Holthuis, L. B., 1971. The Atlantic species of the deep- sea genus Glyphocrangon A. Milne Edwards, 1881. Biological results of the University of Mi- ami Deep-Sea Expeditions, 75.—Bulletin of Marine Science 21(1):267—373. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):929-935. 1998. A new stomatopod (Crustacea: Malacostraca) of the genus Harpiosquilla Holthuis, 1964 from Taiwan and Australia Shane T. Ahyong, Tin-Yam Chan, and Y. J. Laio (STA) Department of Marine Invertebrates, Australian Museum, 6 College St, Sydney South, NSW 2000, and School of Biological Sciences, University of New South Wales, Sydney 2052, Australia; (I-YC, Y-JL) Institute of Marine Biology, National Taiwan Ocean University, Keelung, Taiwan, R.O.C. Abstract.—A new stomatopod, Harpiosquilla ocellata, is described from Tai- wan and Australia. Harpiosquilla ocellata is closely related to H. annandalei (Kemp, 1911), but differs in bearing unarmed submedian carinae on the fifth abdominal somite and in reaching a much larger size. The two species closely resemble each other in color pattern, but differ in the markings of the telson and uropods. The genus Harpiosquilla Holthuis, 1964, includes large squilloids bearing erect spines on the opposable margin of the pro- podus of the raptorial claw, and a deeply excavate posterolateral margin of the cara- pace. All species of Harpiosquilla are re- stricted to the tropical Indo-West Pacific and are associated with soft, level sub- strates. Manning (1995) recognized 11 spe- cies of Harpiosquilla and provided a key to the species. As part of ongoing study of the stomatopod faunas of Taiwan and Australia, we independently discovered a new species of Harpiosquilla from these two regions, which is described below. Specimens are deposited at the National Taiwan Ocean University, Keelung (NTOU), The Taiwan Museum, Taipei (TMCS) and the Australian Museum, Syd- ney (AM). All measurements are in milli- meters (mm). Terminology and size de- scriptors generally follow the conventions of Manning (1969b, 1977), supplemented by some abbreviations proposed by Maka- rov (1979). Total length (TL) is measured along the midline between the apex of the rostral plate and the apices of the subme- dian teeth of the telson. Carapace length (CL) is measured along the midline and ex- cludes the rostral plate. Corneal index (CI) is given as 100CL/cornea width. Uropod segments are measured dorsally, along the midline. Dorsal carinae are abbreviated as follows: median (MD); submedian (SM); intermediate (IM); lateral (LT); and margin- al (MG). The following abbreviations are also used: collector (coll.); antennule (A1), antenna (A2), abdominal somite (AS); tho- racic somite (TS); maxilliped (MXP); Fish- eries Research Vessel (FRV). Systematic Account Harpiosquillidae Manning, 1980 Harpiosquilla Holthuis, 1964 Harpiosquilla ocellata, new species Figs. la—g, 2a—b. Harpiosquilla annandalei.—Graham et al. 1993a:69; 1993b:73 [not H. annandalei (Kemp, 1911)]. Material examined.—Holotype. Taiwan, southwestern coast, Tong-Kong, Ping-Tong County, on sandy mud substrate, depth about 200 m, commercial trawler, 2 Dec 1995, male TL 157 mm (NTOU H-1995- 12-2). Paratypes. Taiwan: on sandy-mud sub- strate, depth about 200 m, commercial 930 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. a-g: Harpiosquilla ocellata, new species: a—e, holotype male, TL 157 mm, Taiwan, (NTOU H-1995- 12-2); f. male, TL 159 mm, Australia (AM P41823); g. paratype male TL 200 mm, Taiwan (AM P51185). h— j: H. annandalei (Kemp. 1911): h-i, male, TL 111 mm. Taiwan (AM); j. male, TL 128 mm, Taiwan (NTOU). a. anterior cephalon; b, i TS5—8, right dorsal; c. telson, left lateral view; d, j. posterior abdominal somites and telson e. uropodal protopod, left ventral view; f. TSS—7 right dorsal; g, h. TS8 sternal keel, right lateral view. VOLUME 111, NUMBER 4 Fig. 2. a—b: Harpiosquilla ocellata, new species, male paratype, Taiwan; c—d: H. annandalei (Kemp, 1911), Taiwan. a, c. dorsal view; b, d. posterior abdominal somites and telson. 932 trawlers. Northeastern coast, Su-Aou, [-Lan County, 16 May 1991, 1 female TL 198 mm (TMCS-0103); 9 Nov 1995, 1 male TL 154 mm, 1 female TL 195 mm (NTOU P- 1995-11-9), 1 male TL 155 mm (AM P51184).—Northeastern coast, Ta-Shi, I- Lan County, 25 May 1998, 1 female TL 176 mm (AM P53156).—Southwestern coast, Tong-Kong, Ping-Tong County, 26 Jan 1994, 1 female TL 232 mm (NTOU P- 1994-1-26); 2 Dec 1994, 2 males TL 157-— 180 mm (NTOU P-1995-12-2); 5 Aug 1996, 1 male TL 197 mm (NTOU P-1996- 8-5), 1 male TL 200 mm (AM P51185). Other material. Australia: East of Port Hunter, Newcastle New South Wales, FRV Kapala, coll. K. Graham, [32°55’S, 151°57’], depth 72 m, 13 Apr 1992, 1 male TL 145 mm (AM P41785); [32°55’S, 151°57’], depth 65-72 m, 2 Nov 1995, 1 female TL 148 mm (AM P49681); [32°54’S, 151°59’E], depth 73 m, 5 Sep 1991, 1 male TL 159 mm (AM P41823).—East of Swains Reef, Queensland, commercial trawler, coll. J. K. Lowry & K. Dempsey [22°28.34’S, 152°59.45’E to 22°26.75'S, 153°09.17’E], depth 137 m, 8-9 Sep 1995, 4 females TL 171—206 mm (AM P49682-49685). Diagnosis.—A1 peduncle longer than CL, but shorter than CL and rostral plate combined. Rostral plate apex broadly rounded, lacking anterior projection. Cara- pace with MD carina. Raptorial claw dac- tylus with 8 teeth. TS5 intermediate carina produced to a single short spine, directed laterally. TS6—8 (rarely only 7—8) with dis- tinct SM and armed IM carinae. TS8 sternal keel broad with angular apex, inclined pos- teriorly. ASI—6 SM carinae distinct. Ab- dominal carinae spined posteriorly as fol- lows: SM 6; IM 1-6; LT 1-6; MG 1-5. Telson MG carina approximately twice length of carina of lateral tooth; MD carina with pair of large, dark, white-margined “eye spots’’ proximally. Exopod of uropod with distal segment black on inner half only. Description.—TL of adults to 232 mm. Dorsal surface lightly pitted. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Eye large, cornea strongly bilobed; CI 278-362 (294-311 in Taiwanese material, 278-336 in Australian material). Ophthal- mic somite with anterior margin broadly rounded. Al peduncle 0.99—1.07 CL, but shorter than CL and rostral plate combined. Al so- mite with slender dorsal processes, with acute apices, directed anterolaterally. A2 scale length 0.71—0.80 CL. Rostral plate slightly broader than long; apex broadly rounded; margins convex; lacking anterior projection or dorsal carina. Carapace anterior width 0.40—0.45 CL; anterolateral spines not extending to base of rostral plate; with MD, IM, LT and MG ca- rinae; MD carina distinct, lacking branches of anterior bifurcation; posterior margin medially concave. Raptorial claw dactylus with 8 teeth; out- er margin broadly curved, slightly angular in adult males; propodus opposable margin with 1—2 smaller spines and several minute denticles between largest spines. Mandibular palp 3-segmented. MXP1—5 with subcircular epipod. MXP5 basal seg- ment unarmed. Pereiopods 1—3 basal segment unarmed; endopod 2-segmented, distal segment sty- liform. TS6-—8 (rarely only 7—8) with distinct, di- vergent SM and armed IM carinae. TS5 intermediate carina produced to a single short spine, spinular or triangular, di- rected laterally; ventral spine triangular, di- rected anteroventrally. TS6-—7 lateral process bilobed; anterior lobe very small and low, apex blunt; pos- terior lobe broad and triangular, apex acute or secondarily bifurcate. TS8 anterolateral angle triangular; sternal keel broad with angular apex, inclined pos- teriorly. AS1-—6 with normal complement of ca- rinae; SM carinae distinct, subparallel or faintly divergent. AS6 posterior margin faintly crenulate adjacent to submedian spines; with sharp ventrolateral spine ante- rior to uropodal articulation. Abdominal ca- VOLUME 111, NUMBER 4 rinae spined as follows: SM 6; IM 1-6; LT 1—6; MG 1-5. Telson longer than broad; submedian, in- termediate and lateral teeth slender, with tu- berculate dorsal carinae and apices deflect- ed dorsally. MG carina approximately twice length of carina of lateral tooth. Submedian and intermediate denticles triangular; some apices spiniform; lateral denticle rounded. Denticles: 5—9, 9-15, 1. MD carina high, uninterrupted proximally; posteriorly armed with short apical spine overhanging several blunt tubercles; with pair of large, dark, white-margined “‘eye spots’ proxiinally. Telson dorsolateral surface rugose, with curved rows of very shallow pits. Telson ventral surface with tuberculate postanal ca- rina, extending half distance between anus and posterior margin. Uropod protopod with small, flattened, ventral lobe anterior to endopod articula- tion; inner margin crenulate. Terminal spines of uropod protopod with blunt lobe on outer margin of inner spine, margin con- cave. Uropod exopod proximal segment outer margin with 8—9 (usually 9) graded movable spines, distalmost longest, not ex- ceeding midlength of distal segment; distal margin with stout ventral spine. Uropod ex- opod distal segment slightly longer than proximal segment; black on inner half only. Color in life.-—(Fig. 2a, b) Eye with cor- nea metallic green. Overall dorsal color light golden brown. Second and third segments of A1 peduncle with one proximal and one dis- tal black spot. Black transverse bar present between Al and ophthalmic somites. Cara- pace with anterolateral and posterior mar- gins, carinae and grooves outlined in dark pigment. Merus of raptorial claw with inner distal black spot and yellow meral depres- sion. Propodus of raptorial claw yellow dis- tally. Posterior margin of thoracic and ab- dominal somites black. AS2 with black me- dian transverse bar. ASI and AS3-—5 with traces of broken transverse bar. SM carinae of thoracic and abdominal somites pale pur- ple. Telson with pair of large, dark, white- margined, “eye spots” proximally; MD ca- 933 rina and carinae of marginal teeth maroon. Spines of uropodal protopod pinkish. Uro- podal endopod yellowish, blackish distally. Uropodal exopod distal segment yellow on outer half and black on inner half. Size.—Taiwanese material: males (n = 7) TL 154—200 mm; females (n = 4) TL 176— 232 mm. Australian material: males (n = 2) TL 145-159 mm; females (n = 5) TL 148— 206 mm. Etymology.—Latin, ocellata, marked with spots, from ocellus, diminutive of oc- ulus, eye, in allusion to the large, eye-like pigment spots of the telson. Remarks.—With the exception of Har- piosquilla annandalei (Kemp, 1911), H. ocellata can be immediately distinguished from all known species of the genus by the combination of the short, apically rounded rostral plate and armed IM carinae of the exposed thoracic somites (Fig. 1b, f, i). This new species closely resembles H. annandalei in most characters, including coloration. Comparison of H. ocellata with the accounts of H. annandalei in Manning (1969a, 1995) and 20 Taiwanese specimens in the collections of the NTOU (2 speci- mens transferred to AM) show the follow- ing differences between the two species: in H. ocellata the SM carinae of AS5 are un- armed (Fig. 1d) whereas they are always armed in H. annandalei (Fig. 1j); in H. ocellata the TS8 sternal keel is broad with an angular apex (Fig. 1g) whereas in H. an- nandalei the keel is hooked (Fig. 1h); A. ocellata attains a much larger size (TL to 232 mm) than H. annandalei (TL to 150 mm (Manning, 1969a), and to TL 130 mm in the 20 Taiwanese specimens examined). Although coloration of the two species is very similar, the body of H. annandalei is generally lighter, the margins of the “eye spots”’ on the telson of both the white and black circles are more sharply defined (Fig. 2c, d) and the MD carina of the telson is colored like the rest of the telson. In con- trast, the body of H. ocellata is light golden brown, the margins of the “eye spots” are somewhat diffuse, and the MD carina of the 934 telson is purplish (Fig. 2a, b). In addition, in H. annandalei the distal segment of the uropodal exopod is blackish with a yellow- ish madrib (Fig. 2d), whereas it is blackish on the inner half and yellowish on the outer half in H. ocellata (Fig. 2b). Although, H. ocellata could easily be confused with H. annandalei, the aforementioned differences will readily separate the two species. Secondary sexual dimorphism is much less evident in H. ocellata than most other species of the genus. As in AH. annandalei, the margins of the telson and MD carina are not inflated, the outer margin of the dacty- lus of the raptorial claw is faintly sinuous and the bases of the dactylar teeth are only slightly inflated. Meristic counts and corneal indices over- lap between Taiwanese and Australian mate- rial. Only minor morphological differences were noted between Taiwanese and Austra- lian material of H. ocellata. Thus, the apices of the posterior lobes of the lateral processes of TS6—7 usually terminate in an acute spine, but are secondarily bifurcate in two speci- mens from Australia, resembling the condi- tion reported by Manning (1969a) for some specimens of H. annandalei. In all Australian specimens, the IM carinae of TS6—8 are armed posteriorly, whereas in three of the 11 Taiwanese specimens, the IM carinae of TS6 are unarmed. Nevertheless, TS7—8 are armed in all Taiwanese specimens. The precise habitat of H. ocellata is un- known although all material was taken on sand and mud substrates with benthic trawls. Harpiosquilla ocellata is presently known only from Taiwan and eastern Aus- tralia in relatively deep water. This, appar- ently disjunct distribution, likely reflects sampling effort but may also result from confusion of H. ocellata with H. annandal- ei by previous workers, especially in pre- served material lacking pigmentation. Fur- ther, the bathymetric range of H. ocellata (63 to about 200 m) coincides with that re- ported by Manning (1969a) for H. annan- dalei (15—206 m). Distribution.—Western Pacific. Known PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with certainty from Taiwan (eastern and southern coasts) and eastern Australia (cen- tral Queensland south to the Newcastle Bight, New South Wales) at depths of 63 to about 200 m, on soft substrata. Acknowledgments We sincerely thank Dr R. B. Manning (National Museum of Natural History, Smithsonian Institution, Washington, D.C.) for kindly providing us with many valuable suggestions and comments on the status of this new species. Thanks also to Drs R. B. Manning and R. Lemaitre (National Muse- um of Natural History, Smithsonian Institu- tion, Washington, D.C.) for reviewing the manuscript and Mr K. Graham (NSW Fish- eries) for providing specimens for study. Dr G. D. Wilson (Australian Museum) offered numerous useful comments on an earlier draft. This study has been partially support- ed by an Australian Postgraduate Award to STA, administered by the University of New South Wales, and a research grant to the TYC from the National Science Council (NSC 88-2313-B-019-027), Taiwan, R.O.C. Literature Cited Graham, K. J., G. W. Liggins, J. Wildforster, & S. J. Kennelly. 1993a. Kapala Cruise Report No. 110. New South Wales Fisheries. ; , & . 1993b. Kapala Cruise Report No. 112. New South Wales Fish- eries. Holthuis, L. B. 1964. Preliminary note on two new genera of Stomatopoda.—Crustaceana 7(2): 140-141. Kemp, S. 1911. Preliminary descriptions of new spe- cies and varieties of Crustacea Stomatopoda in the Indian Museum.—Records of the Indian Museum 6(2):93—100. Makarov, R. R. 1979. A collection of stomatopod crus- taceans of the genus Clorida Eydoux & Souley- et, 1842, from Tonkin Bay, Vietnam.—Crusta- ceana 37(1):39—56. Manning, R. B. 1969a. A review of the genus Har- piosquilla (Crustacea, Stomatopoda), with de- scriptions of three new species.—Smithsonian Contributions to Zoology 36:1—41. . 1969b. Stomatopod Crustacea of the western Atlantic. Studies in Tropical Oceanography 8: VOLUME 111, NUMBER 4 935 viii + 380 pp. University of Miami Press, Mi- of Recent stomatopod Crustacea, with diagno- ami. ses of six new families.—Proceedings of the Bi- . 1977. A monograph of the West African sto- ological Society of Washington 93(2):362-372. matopod Crustacea.—Atlantide Report 12:25— . 1995. Stomatopod Crustacea of Vietnam: the 181. legacy of Raoul Seréne.—Crustacean Research, . 1980. The superfamilies, families, and genera Special Number 4:339 pp. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):936—-941. 1998. A new species of the genus Bellator (Pisces: Triglidae), with comments on the trigloids of the Galapagos Islands William J. Richards and John E. McCosker (W.J.R.) NOAA Fisheries, Southeast Fisheries Science Center, 75 Virginia Beach Drive, Miami, Florida 33149, U.S.A.; (J.E.McC.) California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. Abstract.—A new, endemic species, Bellator farrago, is described from the Galapagos Islands. It differs from its congeners by a combination of characters including absence of a supplemental preopercular spine, short first dorsal fin spine, produced rostral spines, and lack of scales on the breast and interpelvic area. The Galapagos trigloid fauna comprises: B. farrago, Prionotus miles, Pr. stephanophrys, and Peristedion crustosum. The fishes of the New World genus Bel- lator were revised by Miller & Richards (1991); they treated four Atlantic species and three eastern Pacific species. Recent collecting in and around the Galapagos by the submersible Johnson Sea-Link revealed the presence of an undescribed species. The submersible captured one specimen and an- other was found in the California Academy of Sciences Fish Collection. Prior to this study only one Bellator was known from the Galapagos based on Gruchy (1970), who added Prionotus loxias (=Bellator lox- las) to the Galapagos fauna. A re-exami- nation of the Gruchy specimens reveals that they also are this undescribed species and not B. loxias. This new species is diagnosed and described, and details of its habitat are provided based on observations made from the submersible. The submersible also cap- tured three specimens of Peristedion crus- tosum Garman which is a new record of this species from the Galapagos. Methods.—Counts and measurements follow Miller & Richards (1991). Bellator farrago, new species (Figs. 1-3, Tables 1-3) [non] Prionotus loxias: Gruchy 1970:526 (misidentification). Miller & Richards 1991:646 (distribution, following Gruchy 1970). Bussing 1995:1646 (distribution). Material examined.—Holotype: CAS 54562 (103.9 mm SL), southeastern Pacific, Ecuador, Galapagos Islands, Isla Santa Cruz, Academy Bay, R/V Te Vega Field No. TV24-VIII-68. 24 Aug 1968. Para- types: CAS 86564 (96.2 mm SL), south- eastern Pacific Ocean, Ecuador, Galapagos Islands, Isla Genovesa (Tower). 0°21.8'S, 89°58.2'W, JSL dive 3974, 462 m, coll. J. E. McCosker et al., 24 Nov 1995. NMC 69- 78, 2 (75.5-112.4 mm SL), southeastern Pacific Ocean, Ecuador, Galapagos Islands, Isla Isabella, mouth of Tagus Cove, coll. Barr et al. 8 Mar 1968. Diagnosis.—A species of Bellator with opercular spine short, breast and interpelvic area lacking scales, belly scaled, first dorsal spine shorter than second, supplementary preopercular spine weak or absent, anal rays 11, pored lateral line scales 50—52. Description.—Morphometric data are in Table 1, meristic data in Table 2, and com- parison with other eastern Pacific Bellator in Table 3. Head moderate with produced rostrum with small spines on distal edge; small spines on lateral edge of first, second, and third infraorbital bones; no nasal spine, VOLUME 111, NUMBER 4 Fig. 1. SL. Illustration by Molly Brown. although all head bones with very small spines giving rough texture; prominent spines on anterior and posterior edge of or- bit; prominent nuchal, parietal, opercular and preopercular spines, but supplemental preopercular spine absent in three speci- mens and very weakly developed in one. Cleithral spine short, not prominent. First spine of the first dorsal fin shorter than sec- ond spine, serrate anteriorly. Base of first ray of second dorsal fin with serrate anterior edge. Teeth present on premaxillaries, den- taries, head of vomer, extending posteriorly from head of vomer on palatines. Nape, prepectoral area, opercle, breast, interpelvic area scaleless. Belly with scales. Lateral line with 50—52 scales bearing pores, body scales small, with cteni, with 10 rows above the lateral line and 24—34 rows below. Gill rakers on first arch include small rudiments on epibranchial and hypobranchial ‘with short rakers on epibranchial and ceratobran- chial (Table 2). First dorsal, second dorsal and anal fins with 11 elements each. Pec- toral fin with 12—13 connected rays, 3 free rays. (Connected pectoral rays difficult to count.) Vertebrae 26 (9 + 17 on holotype 937 Lateral view and dorsal view of head of Bellator farrago, new species, paratype CAS 86564, 96 mm and paratype CAS 86564). Swimbladder with extrinsic and intrinsic musculature. One papilla present on each eye at one o’clock on the following specimens: CAS 86564, NMC 69-78 (right eye only of 112.4 mm SL specimen). No papillae on eyes of holotype. Mandible with flat ridge about % its length. Coloration.—Two prominent dark sad- dles below each dorsal fin and less distinct dark area on dorsal surface of caudal pe- duncle. Small, irregular, dark spots on dor- sal surface of head. Dark areas on margin of interspinous membrane of first dorsal fin; dark margin on anterior edge of membrane of second dorsal fin. Trunk area below lat- eral line, and anal and caudal fins lack any dark areas, being basically pale. Pectoral fin with middle rays dark from base to tip. From color photograph of a specimen ap- proximately 1 hour after death (Fig. 2): specimen bright red with dorsal saddles ap- pearing as deep red, margin of first few in- terspinous membranes of first dorsal black, as well as ventral edge of caudal peduncle extending onto lower caudal rays. Dorsal fins and anal fin red with scattered yellow 938 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 2-3. 2, Photograph of a paratype of Bellator farrago, new species (CAS 86564), taken approximately one hour after death. 3, Photograph of Peristedion crustosum taken form submersible Johnson Sea-Link at 486 m, seamount SE of Isla San Cristobal, Galapagos. spots on second dorsal fin. Dark spots not visible on head but faint indication of yel- low spots. Medial rays of caudal also yel- lowish. Pectoral fin with black medial rays extending to tip. Lower flank and bottom of head white. Etymology.—From the Latin farrago, a medley or mixture, in reference to the new species’ combination of its congeners’ char- acters, here considered a noun in apposi- tion. Discussion.—This new species is endem- VOLUME 111, NUMBER 4 939 Table 1—Morphometric characters of Bellator farrago, new species. Specimens CAS 54562 CAS 54564 NMC 69-78 NMC 69-78 Character (mm) %SL (mm) %SL (mm) %SL (mm) %SL Standard length 103.9 96.2 112.5 Wd Head length 37.5 36.1 33.7 35.0 38.8 34.5 24.8 32.8 Rostral length 3.1 3.0 2.8 29) 1.9 1.7 2.1 2.8 Rostral width 4.0 3.8 3.9 4.1 3.6 BP Bel 4.1 Snout length 16.5 15.9 15.8 16.4 14.3 12.7 9.3 12.3 Premaxillary length 15.5 14.9 12.7 13.2 15.3 13.6 9.8 13.0 Orbit length 10.4 10.0 8.3 8.6 11.3 10.0 7.0 9.3 Orbit depth 10.9 10.5 7.5 7.8 11.6 10.3 6.0 V2 Interorbital width 3.9 3.8 3.8 4.0 4.9 4.4 3.8 5.0 Cheek height 8.7 8.4 8.5 8.8 8.9 V2) 5.6 7.4 Opercular spine length 10.1 97 98 10.2 11.3 10.0 V2 SES) Preopercular spine length 9.9 9.5 9.0 9.4 9.2 8.2 VP 9.5 Cleithral spine length 8.3 8.0 TI) 8.2 7.6 6.8 7.0 9.3 1st Dorsal spine length 8.3 8.0 8.2 8.5 10.4 9.2 8.0 10.6 2nd Dorsal spine length 13.3 12.8 12.2 12.7 13.2 11.7 10.5 13.9 3rd Dorsal spine length 14.4 13.9 13.8 14.3 17.3 15.4 11.2 14.8 2nd Dorsal fin base length DSS) 24.5 25.7 26.7 28.4 D2 18.5 24.5 Anal fin base length 30.9 29.7 29.3 30.5 33.2 29.5 22.6 29.9 Pectoral fin length 34.7 33.4 31.8 33.1 34.5 30.7 24.2 32.1 1st Free pectoral ray length 32.8 31.6 24.2 25.2 32.5 28.9 19.1 25.3 2nd Free pectoral ray length 28.1 27.0 21.5 Db3) 28.6 25.4 16.9 22.4 3rd Free pectoral ray length 25.4 24.4 19.0 19.8 24.5 21.8 13.2 W/E) Pelvic fin length 30.6 29.5 28.1 29.2 BVP) 28.6 21.9 29.0 Body depth 28.9 27.8 23.6 24.5 27.2 24.2 19.1 DS Least depth of caudal peduncle 7.3 7.0 7.1 7.4 8.4 VD) 6.1 8.1 Gill raker length 2.8 2.6 2.4 2.5 4.4 3.9 1.8 2.4 Snout angle (degrees) 43.0 28.0 34.0 33.0 Table 2.—Meristic characters of Bellator farrago new species. Specimens Character CAS 54562 CAS 54564 NMC 69-78 NMC 69-78 Std. length (mm) 103.9 96.2 112.5 WS 1st Dorsal fin rays 11 2nd Dorsal fin rays 11 Anal rays 11 Pectoral fin rays 13+3 12+3 12+3 12+3 Gill rakers Epibranchials 1+1r 1+I1r Ceratobranchials 7+3r 10+I1r 10 Hypobranchials Total gill rakers 11 Squamation Lateral line scales 52 50 Scale rows above lateral line 10 Scale rows below lateral line 34 24 940 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Characters for distinguishing the eastern Pacific species of Bellator. Characters gymnostethus Supplemental preopercular spine _— present First dorsal spine shorter than 2nd Breast & interpelvic area naked Rostral spines produced ic to the Galapagos Islands and is easily distinguished from its congeners. Its pos- sesses character states present in other Pa- cific Bellator (see Table 3); however, in combination it differs from each. Its color- ation is most like that of B. loxias Jordan (in Gilbert 1896) and it has a similar but notably different preopercular spine condi- tion, but differs in squamation and rostral spine condition. The new species’ reduced first anterior dorsal spine is like that of B. gymnostethus Gilbert 1891, from which if differs in its coloration and preopercular spine condition. Bellator farrago is similar to B. xenisma (Jordan & Bollman, 1890) in coloration and in its rostral spine condition, but differs in its preopercular spine state, its first dorsal spine condition, and in lacking scales on the breast. Its exact relationship to other species of Bellator cannot be de- termined at this time as character polarity for the genus has not been developed nor have cladistic analyses been made. It is pre- mature to speculate further. A paratype of the new species (CAS 86564) was collected by the junior author while aboard the submersible (JSL). It was captured using the submersible’s suction device and photographed at the surface soon after its death (Fig. 2). The water depth at the collection site was 462 m, tem- perature 8.58°C, and the bottom was largely black volcanic rock overlain with sand patches and small boulders partially en- crusted with a thin cover of stony corals and sponges. As a result of our studies, we conclude that the Galapagos trigloid fauna comprises: Bellator farrago, Prionotus miles, and Pr. stephanophrys of the family Triglidae and Species loxias xenisma farrago absent present absent longer than 2nd longer than 2nd _ shorter than 2nd scaled scaled naked not produced produced produced Peristedion crustosum Garman, 1899 of the Family Peristediidae. As stated in the intro- duction, the previous record of B. loxias (as Prionotus loxias, Gruchy 1970) from the Galapagos was based on material which we describe herein as B. farrago. Bellator lox- ias reaches offshore to the Cocos Island area as noted by Bussing (1995) and a spec- imen collected by the Albatross (MCZ 30779, 81.0 mm SL) which was identified by the senior author. The species of Prio- notus reported from the Galapagos also de- serve explanation. Prionotus miles Jenyns, 1842 is an endemic species and has been captured at several locations throughout the islands. Galapagos records of P. albirostris Jordan & Bollman (1890) (Gruchy 1970, Bussing 1995, Grove & Lavenberg 1997) are based on misidentifications of P. ste- phanophrys Lockington (1881), as con- firmed by the senior author after examina- tion of Gruchy’s specimens. Other errone- ous records of Galapagos triglids are ex- plained by Gruchy (1970:526): including Jordan & Evermann’s (1896:2172) listing of P. horrens Richardson (1845) from Gal- apagos and Teague’s (1951:26) mistake in listing the type locality of P. quiescens Jor- dan & Bollman (1890) as the Galapagos Is- lands rather than the Gulf of Panama. Grove & Lavenberg (1997:302-—303) re- ported upon a poorly preserved specimen (LACM 20838, 138.5 mm SL) of Periste- dion crustosum Garman 1899 taken off Daphne Minor, Galapagos, in 1938 by the Allan Hancock Expedition. They suggested that “‘Peristedion crustosum may prove to be a mainland species, which would make the population in the Galapagos a new spe- cies.” Four specimens of Peristedion were VOLUME 111, NUMBER 4 collected by the junior author on JSL dives and after comparison with the type material of the two eastern Pacific Peristedion by the senior author, we find that the JSL speci- mens are P. crustosum. The specimens are: CAS 86565, 151.6 mm SL, Galapagos Is- lands, Isla Genovesa (Tower Island), 0°21.8'S, 89°58.2’W, JSL Dive 3974, sand bottom strewn with small volcanic rocks, 372 m; and CAS 86562, 103.1 and 108.8 mm SL, Galdpagos Islands, 1°06’S, 89°12.2'W, seamount SE of Isla San Cris- tobal, JSL Dive 3934, volcanic rock and boulder bottom with sand channels, 486 m (Fig. 3). Other specimens were observed at Isla Fernandina, off Cabo Douglas (ISL Dive 3957, 0°17.5'S, 91°38.9'W), at 431 m resting on a 60° sand slope, and off Isla Floreana (JSL Dive 3944, 1°13’S, 90°23’W), above a flat sand bottom at 305 m, the sin- gle specimen collected and deposited at the Instituto Nacional de Pesca in Quayaquil. Collection records for both species of Peristedion are unknown other than the ma- terial upon which Garman based his origi- nal description. Peristedion crustosum was illustrated by Garman in color, but the other species, P. barbiger, was not. The two spe- cies can be easily separated by the nature of the chin and lip barbels and rostral spine shape. In P. barbiger the barbels are un- usually thick and closely spaced giving a thick bearded appearance. In P. crustosum the barbels are slimmer and fewer in num- ber. Peristedion crustosum has a well de- veloped nasal spine but this spine is weak or absent in P. barbiger. The striking dif- ference is the form of the rostral spines. In P. crustosum these spines project forward nearly parallel, wherein P. barbiger these spines project forward in a convergent di- rection, though their tips do not touch. There were indications of thickening of these bones which possibly could be hy- perostosis, a condition found in a few trig- loid fishes. 941 Acknowledgments The late André de Roi generously donat- ed the holotype of the new species. We thank R. Grant Gilmore and the staff of Harbor Branch Oceanographic Institution for the use of the submersible Johnson Sea- Link. For assistance and permission to study in Ecuador, we thank: Oscar Aguirre, Sub- director de Pesca de Galapagos; Harold Miller, Franklin Ormaza-Gonzdlez, and Carlos Villon, Instituto Nacional de Pesca; Arturo Izurieta Valery and Eduardo Ama- dor, Parque Nacional Galapagos; Alejandro Villacis, Captain de Puerto de Isidro Ayora; and Chantal Blanton, Director de Estacié6n Cientifica Charles Darwin. Sylvia Lafram- boise (NMC) kindly loaned us the Gruchy (1970) specimens and Karsten Hartel (MCZ) kindly loaned us the Peristedion types and B. loxias. Molly Brown prepared the illustration. We also thank the David and Lucile Packard Foundation and the Dis- covery Channel for grants and other assis- tance. Literature Cited Bussing, W. A. 1995. Triglidae. Pp. 1643-1648 in W. Fischer, E Krupp, W. Schneider, K. E. Carpen- ter, & V. H. Niem, eds., C. Pacifico Centro-Ori- ental, Guia FAO para la identificacion de es- pecies para los fines de la pesca. Sommer. 3(2): 1201-1813. [In Spanish] Garman, S. 1899. The fishes. Jn Reports on an explo- ration off the west coasts of Mexico, Central and South America, and off the Galapagos Is- lands, in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer “Albatross,” during 1891, Lieut. Commander Z. L. Tanner, U.S.N., commanding. Memoires of the Museum of Comparative Zoology. 24: text: 1-431, Atlas: Pls. 1-97. Grove, J. S., & R. J. Lavenberg. 1997. The fishes of the Galapagos Islands. Stanford University Press, Stanford, California. 863 pp. Gruchy, C. G. 1970. A second Prionotus birostratus Richardson, with notes on the distribution of Prionotus in the southeastern Pacific Ocean— Pacific Science 24:523—527. Miller, G. C., & W. J. Richards. 1991. Revision of the western Atlantic and eastern Pacific genus Bel- lator (Pisces: Triglidae).—Bulletin of Marine Science 48:635—656. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):942—953. 1998. A new species of Polyipnus (Teleostei: Stomiiformes) from the western Indian Ocean, with comments on sternoptychid ecology Antony S. Harold, James H. Wessel, III, and Robert K. Johnson Grice Marine Laboratory, Department of Biology, College/University of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412, U.S.A. Abstract.—Polyipnus limatulus is described based on material collected in the Gulf of Aden and immediately adjacent areas of the northwestern Indian Ocean. These collections were made during midwater fish surveys conducted from three vessels, R/V Beinta (1987), R/V Malcolm Baldridge (1995), and R/V Tyro (1992). Like other species in this genus P. limatulus appears to occur in association with the continental slope and/or various types of sea floor rises. Polyipnus limatulus shares certain apomorphic characters with the P. spinosus species group; among these are a multispinose posttemporal, and the occurrence of numerous spine-like denticles on the modified scales covering many of the ventral photophores. These denticles are lacking on the ACB photophore scales, a character which distinguishes the new species from a putatively closely re- lated species, P. asper (eastern Indian ocean). Polyipnus indicus (western In- dian Ocean) has somewhat similar scale denticulation but has a relatively un- modified parietal, among other differences. The new species was found to occur in relatively warm, saline, low oxygen water associated with Red Sea outflow, whereas the other Polyipnus species collected during the R/V Tyro survey, P. omphus Baird, 1971 was collected in the colder, less saline, higher oxygen water associated with the Somali Current. The Stomiiformes are mainly meso- and bathypelagic fishes, found in all oceans but more diverse in the tropics and subtropics (Harold & Weitzman 1996, Harold 1998). Among the Sternoptychi- dae or hatchetfishes one genus in partic- ular, Polyipnus, contains more species than any of the other nine genera in the family (Weitzman 1974, Harold 1994). Polyipnus has been the subject of several revisionary studies (Schultz 1938, 1961, 1964; Baird 1971; Borodulina 1979; Har- old 1994) and a series of recent species descriptions (Harold 1989, 1990a; Aizawa 1990; Last & Harold 1994). The most di- verse clade of this genus, the Polyipnus spinosus species group, was phylogeneti- cally diagnosed by Harold (1994:455-— 458) and at that time contained 14 named species. More recently, Last & Harold (1994) described P. latirastrus from the Coral Sea and added it to the group, based on the presence of synapomorphies pro- posed by Harold (1994). We describe here a new species belong- ing to Polyipnus, based on collections made in the Gulf of Aden and vicinity by three vessels, R/V Beinta (January and March 1987), R/V Malcolm Baldridge (August 1995), and R/V Tyro (August 1992 and Jan- uary 1993, as part of the Somali Ecosystem Study [SES], Netherlands Indian Ocean Program [NIOP]). A total of 192 specimens were examined, ranging in standard length from 6.5 to 45.7 mm. For the first time a reasonably complete series of larval through adult specimens is available for a species of the P. spinosus species group and we therefore illustrate and briefly describe the larva. VOLUME 111, NUMBER 4 Materials and Methods Specimen body size is standard length (SL) in all cases. Morphometric, meristic, photophore and other characters were de- termined following the methods and ter- minology of Harold (1994). Values for the holotype are given in parentheses in the de- scription. Terminology of the photophores is repeated here for convenience, and be- cause the terms now in use replace those of Schultz (1961) that occur in much of the sternoptychid literature. The abbreviation for each photophore cluster and its verbal descriptor, as defined by Schultz (1961), are given in parentheses: ACA (SAN, supra- anal), ACB (AN, anal), ACC (SC, subcau- dal), BR (BR, branchiostegal), IP CIS, isth- mus), L (L, lateral), OP (PRO, preopercular + SO, subopercular + PTO, postorbital), ORB (PO, preorbital), OVA (SP, suprapec- toral), OVB (SAB, supra-abdominal), PV (AB, abdominal), and VAV (PAN, preanal). Specimens were cleared and counterstained for bone and cartilage following the meth- ods outlined by Pothoff (1984) and Taylor and Van Dyke (1985). Observations on os- teology and dentition of P. limatulus were based on two cleared and stained adult specimens (USNM 345149, 31.8—38.0 mm SL). Institutional abbreviations are as listed in Leviton et al. (1985). Acronyms used for various ““Subareas”’ of the Somali Ecosystem Study are defined as follows: GAD (Gulf of Aden), between 12°20’ and 12°45'N, 38°08’ and 50°05’E; SWL (Great Swirl, an eddy created by the Somali Current), between 6°10’ and 12°05'N, 53°40’ and 55°00’E; USI (Upper Somali Current Inshore), between 7°30’ and 10°50’N, 50°21’ and 52°05’E; LSI (Lower Somali Current Inshore), between 0°06’ and 3°22'N, 44°56’ and 48°26’E; LSO (Lower Somali Current Offshore), 1°55’N, 53°34’E, represented by a single station. Ecological comparisons were made be- tween P. limatulus and other sternoptychids with which it was collected. A mathemati- cal formula, the “‘weighted hydrographic 943 index’’ (modified, after Wessel 1997) was used to summarize hydrographic conditions with which each species is associated. The analysis is based on 17 hauls of a rectan- gular midwater trawl. Weighted hydrographic index.—To allow comparisons between species, hauls, sta- tions, subareas, day/night occurrence and season, a weighted hydrographic index was devised to best represent the average con- ditions of capture for specimens of a given species under stated capture criteria (e.g., day vs. night; summer vs. winter). Con- struction of this index involved two steps: computation of an integrated average for each hydrographic parameter for each dis- crete depth segment sampled, and combi- nation of this average with proportionate capture information for each species for each set of capture criteria. The hydro- graphic average was constructed as the in- tegral average of the hydrocast data within each sampling depth segment by the meth- od of trapezoidal approximation, evaluated for all hydrographic data points (up to N = 9 discrete values) within a sampling seg- ment (1.e., one “‘haul’’), such that: (ES) where V, = computed integral average for a given hydrographic parameter (temperature, salinity, dissolved oxygen, etc.) for a given haul segment. V..;,V; = two sequentially successive measurements of that value within that segment. V, = the smaller of the ‘two values, V,,, or V;, for each comparison. D,,,,D; = depth at which the two measure- ments, V,,,, V;, respectively, was made; where D;,, > Dj. 944 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. D, = shallowest depth of measure- ment for segment. D, = deepest depth of measurement for segment. Comparisons between species (for given evaluation criteria) were then facilitated by weighting the integral average of the hy- drographic values for each haul (successful in capturing the species) by proportionate capture of the species over all hauls suc- cessful in capturing the species, such that, for species A: INV, = weighted hydrographic index for species A, as restricted by preset criteria (e.g., day vs. night, sum- mer vs. winter). H = number of hauls in which spe- cies A was captured. V,. = integral average of hydrographic value for haul 1. Polyipnus limatulus, holotype, USNM 316316, 39.0 mm SL. n, = number of specimens of species A taken in haul i. Z I total number of specimens of species A taken in all (=H) hauls. Polyipnus limatulus Harold & Wessel, new species Figs. 1-4, Tables 1 & 2 Polyipnus cf. indicus Harold, 1994:437, 450, Sil (USNM 293986530127 7- 316316; fig. 16B, pectoral girdle mor- phology). Type material.—Holotype: USNM 316316, “(', 39:0° mms) Eee 47°57'30"E, (0-)315 m, R/V Beinta, Cr. 18, Sta. 4, 8 Jan 1987. Paratypes: MCZ 150026, “(1,9 28:5" mm) 74 .0s400aNe 50°42'36"E, 0O—1000 m, R/V Malcolm Bald- ridge, Cr. 9506, Sta. 10-50 (1), 9 Aug 1995. MCZ 150036, (1, 27.4 mm), 14°05’00"N, 50°42'36"E, 300-150 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-50 (4), 9 Aug 1995. USNM 293986, (3, 42.6—45.7 mm), 12°01'00'N, 51°16'36’E, (0-)375—393 m, R/V Beinta Cr. 20, Sta. 18, 1600 h, 12 Mar 1987. USNM 301277, (5; 2, 26.4—31.8 VOLUME 111, NUMBER 4 945 Fig. 2. Polyipnus limatulus, larva, MCZ 150024, 6.5 mm SL. 16 14 12 10 ACA & ACB Photophores 0 5 10 15 20 25 30 35 40 45 50 Standard Length (mm) Fig. 3. Plot of ACA + ACB photophore count (combined) versus SL for Polyipnus limatulus. 946 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 50E Fig. 4. Map of northwestern Indian Ocean, off east Africa, showing collection stations of Polyipnus lima- tulus. Polyipnus limatulus: half-filled circle = R/V Beinta stations, half-filled square = R/V Malcolm Baldridge stations, shaded circle = R/V Tyro cruise B1 stations (summer), shaded square = R/V Tyro cruise B2 stations (winter). Polyipnus omphus: open triangle = R/V Tyro cruise B1 stations (summer), open diamond = R/V Tyro cruise B2 stations (winter). mm), 11°57'30"N, 51°07'30’E, (0-)366 m, R/V Beinta, Cr. 20, Sta. 11, 11 Mar 1987. USNM 345149, (56; 14, 32.6—39.9 mm; 2 cleared and stained, 31.8—38.0 mm), col- lected with holotype. ZMA 121.797, (@, 26.7—31.6 mm), 12°07'00"N, 46°54’00’E, 202-300 m, Tyro, Cr. B1, Seq. 582, Sta. GA2, 2304-2353 h, 5 Aug 1992. ZMA 121.798, (2, 25.6—25.7 mm), 12°46’00'N, 46°36'36"E, 200-300 m, Tyro, Cr. B2, Seq. 1129, Sta. GA2, 2134-2244 h, 29 Jan 1993. Non-type larval material: MCZ 150020, (4), 16°46'24’N, 55°27'00"E, 0-1000 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-48 (1); 7- Aug» 1995. MCZ 150021 Gee 16°46'24"N, 55°27'00"E, 500-300 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10—48 (3), 7 Aug 1995. MCZ. 1500227 "GR VOLUME 111, NUMBER 4 947 Table 1.—Morphometric characters expressed as percentages of head length (2 to 6) or standard length (all others) for Polyipnus limatulus holotype (USNM 316316) and 26 paratypes: MCZ 150026(1), 150036(1), USNM 293986(3), 301277(2), 345149(14), ZMA 121.797 (3), and 121.798 (2). Holotype 1. Standard length 39 2. Posttemporal spine length Da 1 3. Preopercular spine length 13.9 4. Head length 36.9 5. Orbit diameter 45.8 6. Snout length 18.8 7. Body depth 61.3 8. Caudal peduncle length 18.7 9. Caudal peduncle depth 10.8 10. Dorsal fin length 17.9 11. Anal fin length DES 12. Preanal length 70.8 13. Predorsal length 59.7 14. Prepelvic length 65.6 15. Postdorsal length 47.7 16. Postanal length 40 17. Dorsal-pelvic length 52.8 18. ACC length (left) 6.4 19. ACC length (right) 0) 20. ACB-ACC length (left) 5.6 21. ACB-ACC length (right) 4.6 16°46'24’N, 55°27'00"E, 300-150 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-48 (4), 7 Aug 1995. MCZ 150023, (6), 16°47'24"N, 55°27'00"E, 0-100 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-49 (1), 7 Aug 1995. MCZ 150024, (1), 16°47'00"N, 55°27'06"E, 500—300 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-49 Cred Aue S995 MEZ 150025, 1): 16°47'00"N, 55°27'06"E, 150—O m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-49 (5), 7 Aug 1995. MCZ 150027, (2), Table 2.—Frequency distribution of the number of ACB photophores in species of Polyipnus with two posttemporal spines. ACB photophores Species 6 1 8 9 10 11 Polyipnus asper 21 8 Polyipnus indicus 1 14 4 Polyipnus limatulus 2 144 9 1 Polyipnus nuttingi 1 22 6 Polyipnus oluolus 1 Paratypes Min Max Mean 25.6 45.7 35 25.7 35.3 30.5 10.1 19.2 15.0 33.7 38.9 35.6 41.1 53.2 46.1 17.4 Ds) 21.0 58.5 69.5 63.3 14.5 18.8 16.5 9.6 12.1 10.8 17.6 21.9 19.5 20.9 24.8 22.9 67.1 72.6 70.4 58.2 64.4 60.9 60.6 69.1 65.1 44 51.9 47.4 36.1 42.7 39.4 51.6 57.3 54.9 5.8 9.2 6.7 5.8 7 6.4 2.4 5.6 3.8 BES) 4.6 4.1 14°03'24"N, 50°54’12”E, 300-150 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-51 (4), 9 Aug 1995. MCZ 150028, (10), 10°40’48"N, 51°51'24"E, 0-900 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-52 (1), 11 Aug 1995. MCZ 150030, (6), 10°40'48"N, 51°51’24’E, 300-150 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-52 (4), 11 Aug 1995. MCZ 150031, (2), 10°42'18"N, 51°49’06"E, 0-900 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-53 (1), 11 Aug 1995. MCZ 150032, (16), 10°42'18"N, 51°49'06"E, 300-150 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-53 (4), 11 Aug 1995. MCZ 150033, (1), 10°45'18"N, 51°52'54"E, 300—150 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-54 (4), 12 Aug 1995. MCZ 150035, (1), 10°45'00"N, 51°52'18"E, 300-150 m, R/V Malcolm Baldridge, Cr. 9506, Sta. 10-55 (4), 13 Aug 1995. MCZ 153593, (6), col- lected with MCZ 150036 (paratype). ZMA 121.799, (1), 07°53’N, 50°37'E, 200-300 m, Tyro, Cr. B2, Sta. US1, 2253-2343 h, 948 18 Jan 1993. ZMA 121.800, (1), 10°57'N, 52°01'E, 200-300 m, Tyro, Cr. B2, Sta. US2, 2121-2204 h, 20 Jan 1993. ZMA 121.801, (2), 12°06'N, 55°01’E, 300-500 m, 7yro, Cr. B2, Sta. SI, 1158-1251 h, 24 Jan 1993. ZMA 121.802, (1), 12°53’N, 50°07'E, 100-300 m, Tyro, Cr. B2, Sta. GA1, 0930-1018 h, 27 Jan 1993. ZMA 121.803, (7), 13°20'N, 50°15’E, 200-300 m, Tyro, Cr. B2, Sta. GA1, 2313-0005 h, 27 Jan 1993. ZMA 121.804, (12), 12°27’N, 47°00’E, 100-300 m, Tyro, Cr. B2, Sta. GA2, 0844-0947 h, 29 Jan 1993. ZMA 121.805, (29), collected with ZMA 121.798 (paratypes). ZMA 121.806, (8), 12°52’'N, 46°31’E, 300-500 m, Tyro, Cr. B2, Sta. GA2, 2331-0030 h, 29 Jan 1993. Diagnosis.—Characters, in combination, which serve to distinguish Polyipnus lima- tulus: Lateral pigment bar tapered and long, approaching or reaching lateral midline. Two posttemporal spines: dorsal spine long, basal spine length less than half of that of dorsal spine. Photophore scale denticles re- stricted to PV, VAV, and ACC clusters; no denticles on ACB photophore scales. ACB photophores 8—10. ACC photophores close- ly juxtaposed, with the separating inter- spaces much less than one half of the width of one ACC photophore. Longitudinal pa- rietal keel or crest terminating posteriorly with a disjunct portion bearing two spines, one directed anteriorly and one directed posteriorly. Autapomorphies: none known. Description.—Moderate-sized species, maximum observed adult body size 45.7 mm SL. D (12) 10-14. A(14)14-17. P(12)11—-13. V(6)5—6. GR(114+11) 10+9- 11+11 = (22) 19-22. Branchiostegal rays 10 (7 anterior ceratohyal + 3 posterior cer- atohyal). Vertebrae (12+21) 12-14+19-2]1 = (33)32-34. Body covered with slightly imbricated, sheet-like, dorsally elongate scales. Other scales thickened and modified in association with photophores (see Harold 1994). Body profile anterior of dorsal fin and pelvic fin broadly ovate, posteriorly ta- pering at a 60° angle to caudal peduncle. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Profile of caudal peduncle slightly elongate, and rectangular. Orbit circular to slightly ovate with elon- gation in dorsoventral axis; pronounced aphakic space located ventrally to lens. Ventral margin of dentary with two keels; lateral-most keel smooth or sparsely serrate, ventromedial keel with 9-15 fine serrae. Premaxillary teeth minute and conical, un- iserial over posterior half of bone and with up to three rows anteriorly; about 55 to 60 teeth in longest series. Maxillary teeth mi- nute and conical, uniserial with 45 to 48 teeth. Dentary teeth minute; conical and un- iserial over posterior half of bone, and slightly recurved in up to four rows ante- riorly; 27 to 33 teeth in longest series. Pal- atine and mesopterygoid teeth absent. Vo- merine teeth minute and conical, in five to seven longitudinal rows arranged in a trans- verse patch across the anterior wing-like process; no teeth present on posteromedian process. Parietal crest discontinuous; ante- rior portion keel-like, posterior region with two conical spines in the medial plane, di- rected anteriorly and posteriorly. Posttem- poral dorsal limbs smooth or ornamented with two or three weak serrae. Two post- temporal spines: dorsal spine without ser- rae, elongate, ranging from 25.7 to 35.3% HL (n = 10), posterodorsally directed; ven- tral spine short, also unornamented (usually about one quarter of the length of the dorsal spine), ventrolaterally directed. Ventral margin of pectoral shield with five to 12 small conical spines, anterior spine slightly deflected laterally. Vertical and anteroven- tral rami of preopercle each with two par- allel, deeply serrate lateral ridges, the num- bers of serrae highly variable. Preopercle with two spines: ventral spine longest, rang- ing from 10.1 to 19.2% HL (n = 10), an- teriorly curved, directed anteroventrally; dorsal spine minute, similar to adjacent ser- rae of adjacent lateral ridge, directed ven- trolaterally. Dorsal spinous process smooth, exposed length and height about equal, ter- minating posterodorsally as two conical dorsolaterally-directed spines (one per VOLUME 111, NUMBER 4 side). Anal-fin pterygiophore spines absent. Morphometric characters summarized in Table 1. Ventral margins of photophore scales smooth, except those of PV, VAV, and ACC clusters which have numerous denticles. OVB 1 + 1 + 1, variably positioned, but most commonly found in an anteriorly-in- clined straight line (in holotype elevation of central OVB slightly ventral to that of first and third OVB). ACA #1 and #2 isolated, #3 joined to ACB forming continuous clus- ter. ACB (9)8—11, usually 9 or 10 (Table 2), with dorsal step between #3 and #4. Color in preservative.—Dark brown pig- ment saddle with an elongate and narrow predorsal notch and a moderately long lat- eral pigment bar tapering toward and usu- ally reaching lateral midline. Dark brown to black chromatophores delineating myosepta and lateral midline. Two prominent, verti- cally elongate patches of dark brown pig- ment at base of caudal fin rays immediately dorsal and ventral to lateral midline, espe- cially distinct in juveniles and larvae. Cen- tral portion of caudal peduncle with internal rectangular patch of dark brown pigment in association with vertical septum. Larval morphology.—Overall larval morphology similar to that described and illustrated by Ahlstrom et al. (1984) for Po- lyipnus polli Schultz, 1961; smallest speci- men of P. limatulus examined (6.5 mm SL; Fig. 2) is slightly deeper bodied in the tho- racic and abdominal regions than the illus- tration of the 5.2 mm specimen of P. polli, in addition to much greater development of the median fin rays and some photophore groups. Evidently, photophore development is relatively protracted in P. polli. For ex- ample in P. polli the first photophores to appear in the SAB (=OVB) cluster do so at a standard length of between 9.0 and 9.6 mm whereas in P. limatulus they are begin- ning to develop at 7.6 mm. Similarly in P. polli the AN (=ACB) photophores begin to appear between 7.5 and 9.0 mm SL whereas in P. limatulus there are two well-devel- oped ACB photophores present on each 949 side in 6.5 and 6.8 mm specimens. In P. limatulus, the combined linear cluster ACA + ACB reaches its full adult complement (11-14) at standard lengths between about 25 and 30 mm (Fig. 3). According to the occurrence of small photophore buds on the posterior margin of the ACB cluster Po- lyipnus polli adds photophores to this clus- ter at a standard length of 23.5 mm so the two species may be similar in the size at which the full complement is attained. These observations are in general agree- ment with other congeners for which pho- tophore development has been reported, ex- cept for P. fraseri (P. spinosus species group) which has paedomorphic develop- ment of the ACB cluster (Harold 1990b). The adults of P. limatulus have two ver- tically elongate patches of dark pigment near the base of the caudal fin rays, as de- scribed and illustrated (Fig. 1). In well-de- veloped larvae and juveniles (9.5 + mm SL) these patches appear as two distinct black spots, one at the base of the ventral caudal-fin rays and one at the base of the dorsal caudal-fin rays, on each side of the fin base. Smaller larvae (6.5—7.6 mm SL) lack the caudal spots (Fig. 2) and any other pigmentation in the caudal region. Etymology.—The specific name limatu- lus is based on the diminutive form of the Latin adjective limatus, meaning filed, pol- ished or smoothed, in reference to the char- acteristic lack of denticles on the scales covering the ACB photophores. Distribution.—Polyipnus limatulus is known to occur only in the Gulf of Aden and immediately adjacent areas in the northwestern Indian Ocean (Fig. 4). Ecology.—Deep-bodied sternoptychids (Sternoptychini, sensu Baird 1986) were represented in the Somali Ecosystem Study (SES) region collections of R/V Tyro (1992, 1993) by 809 specimens allocated among six species: Argyropelecus affinis (n = 336), A. hemigymnus (n = 41), A. sladeni (n = 312), Polyipnus limatulus (n = 66), P. omphus. (n = 21), and Sternoptyx ob- scura (n = 33). Polyipnus limatulus was 950 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A Polyipnus limatulus Cruise 2GAD govt 4USI SLSI Subarea Polyipnus omphus Cruise Fig. 5. Seasonal and spatial distribution of captures (n = number of specimens) of two species of Polyipnus, P. limatulus and P. omphus. Seasonal distribution indicated by B1 (summer) captures and B2 (winter) captures. Spatial distribution indicated by subarea (2GAD = Gulf of Aden, 3SWL = Great Swirl, 4USI = Upper Somali Inshore, 5LSI = Lower Somali Inshore; definition of subareas provided in Materials and Methods). taken at five stations (Fig. 4: GA2, GAI1 (subregion GAD); SI (subregion SWL); US2, US1 (subregion USI)) and in three SES subregions: GAD (n = 62), SWL (n = 2), USI (n = 1). Despite a nearly even distribution of effort among stations by sea- son, geography and depth, P. limatulus was unequally represented in the winter (n = 62) vs. summer (n = 3) cruises. Polyipnus omphus was taken at four stations (Fig. 4: SB2, subregion SWL; US1, subregion UST; USO, SB1, subregion LSI) and in three SES subregions: SWL (n = 1), USI (n = 2) and LSI (n = 18). Polyipnus omphus was sim- ilarly unequally represented in summer (n = 20) vs. winter (n = 1) cruise stations. A comparison of geographic (Figs. 4, 5) and seasonal (Fig. 5) captures of the two Polyipnus species reveals that within the SES area their occurrences are nearly op- posite, with highest captures of P. limatulus in the north and in winter (especially GAD subregion), and highest captures of P. om- phus in the south and in summer (especially LSI subregion). Vertical distributional records are nearly the same in our material, with most speci- mens taken at night (51:15 for P. limatulus, 17:4 for P. omphus) and mostly within the 200 to 300 m depth stratum (68% of all specimens of P. limatulus, 71% of all spec- imens of P. omphus). All captures, day and night, are within the 100 to 500 m range. There is no evidence for diel vertical mi- gration in our data. This conclusion is cor- roborated by the consistency (Fig. 6A, B) for day vs. night values for weighted-inte- gral values of temperature, salinity and dis- solved oxygen (see Wessel & Johnson 1998 and Wessel 1997 for treatment of ““weight- integral’? values for hydrographic data as- sociated with SES net captures of mesope- lagic fishes). With comparison to each other and in comparison with other reported SES ster- noptychids (Fig. 6A, B), P. limatulus and P. omphus are again distributional oppo- sites. P. limatulus is consistently associated with warm, relatively saline, low oxygen water; P. omphus is associated with colder, less saline, higher oxygen water. This prob- ably reflects the influence of Red Sea out- flow water on P. limatulus vs. the summer intrusive occurrence of P. omphus in the SES area, associated with the summertime development of marked coastal and off- shore upwelling in concert with develop- ment of the monsoonal Somali Current flow. Remarks.—The parietal structure of specimens here ascribed to P. limatulus was noted by Harold (1994:511) as an indica- tion that material listed in that study as P. cf. indicus from the Gulf of Aden, the VOLUME 111, NUMBER 4 DAY HAULS: DSALINT vs. DOXYINT vs. DAYINT ANAS NIGHT HAULS: NSALINT vs. NOXYINT vs. NIGHTINT ULROIN So. ab e nmIP e “Bo SP 2: Ss SOs 2S 2 SN Ss 2S = ee SS SF SS © Soe Ss SH sii eos Fig. 6. Comparison of weighted integral-mean val- ues of temperature, salinity, and dissolved oxygen for day (respectively: DAYINT, DSALINT, DOXYINT) and night (respectively: NIGHTINT, NSALINT, NOX- YINT) captures of six species of sternoptychids in the Somali Current Ecosystem Area (see Materials and Methods for explanation of weighted integral-mean values). Open diamond = Argyropelecus affinis, open triangle = A. hemigymnus, filled circle = A. sladeni, cross = Polyipnus limatulus, open square = P. om- phus., filled square = Sternoptyx obscura. 951 northern portion of the geographic range of P. indicus Schultz, 1961, may represent an undescribed species. Further study of ma- terial available at that time, as well as new collections from the region (R/V Malcolm Baldridge and R/V Tyro), has indicated to us that the northern, P. cf. indicus material, should be recognized as a distinct species. Five species of the P. spinosus species group have posttemporal bones with two spines: P. limatulus, P. asper Harold, 1994 (Andaman Sea, eastern Indian Ocean), P. indicus (western Indian Ocean), P. nuttingi Gilbert, 1905 (Hawaiian Islands), and P. ol- uolus Baird, 1971 (Marshall Islands). Po- lyipnus limatulus is most similar to the two other Indian Ocean species, P. indicus and P. asper (couplet 15 of Key to the Species of Polyipnus, in Harold 1994:470—475) in regard to body shape, and fin-ray counts. The new species is more similar to P. asper in regard to the presence of denticles on the ACB photophore scales. Polyipnus limatu- lus and P. indicus, which are both known to occur off east Africa, can be readily dis- tinguished on the basis of the form of the parietal crest. The new species has the bi- spinate parietal morphology as is typical of P. asper and more derived species (see Har- old 1994:436, fig. 6D), whereas in P. indi- cus, like P. nuttingi, and P. oluolus, the pa- rietal crest is a simple blade lacking the posterior ornamentation (Harold 1994:436, fig. 6C). An important feature separating P. limatulus from P. asper is the lack of den- ticles on the ACB photophore scales in P. limatulus. The new species often has more ACB photophores than any of these four other species (Table 2). The other two spe- cies with two posttemporal spines, P. nut- tingi, and P. oluolus, are also readily distin- guished from P. limatulus. Polyipnus nut- tingi, an apparent endemic of the Hawaiian Islands, has relatively high gill raker counts (21—25, usually 23 or 24) compared with P. limatulus (18-21, usually 19). The remain- ing species, P. oluolus (known from a sin- gle specimen from the Marshall Islands; Baird 1971), has an overall photophore 952 configuration, which includes only six ACB, that is unusual for members of the P. spinosus group (Harold 1994:515-517, fig. 52). Phylogenetic relationships of Polyipnus species were proposed by Harold (1994). Ali those species of the P. spinosus group with only two posttemporal spines, of which P. limatulus is one, appear to be bas- al, although not comprising a monophyletic group. Little else can be said of the rela- tionships of the new species until a survey of the characters utilized by Harold (1994) is completed for P. limatulus and P. lati- rastrus. Acknowledgments We thank K. E. Hartel (MCZ), L. Palmer and S. H. Weitzman (USNM), and I. H. Is- bruecker and P. Schalk (ZMA) for making specimens available for this study. Collect- ing efforts in association with R/V Tyro in which JW and RKJ participated were sup- ported through Office of Naval Research grant N 00014-96-11-0469. We thank J. Moore, C. B. Robbins and an anonymous reviewer for their helpful reviews of the manuscript. This paper is Contribution Number 145 to the Grice Marine Labora- tory, College of Charleston, Charleston, South Carolina. Literature Cited Ahlstrom, E. H., W. J. Richards, & S. H. Weitzman. 1984. Families Gonostomatidae, Sternoptychi- dae, and associated stomiiform groups: devel- opment and relationships. Pp. 184—198 in H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fa- hay, A. W. Kendall, Jr, & S. L. Richardson, eds., Ontogeny and systematics of fishes—— American Society of Ichthyologists and Her- petologists, Special Publication Number 1:1— 760. Aizawa, M. 1990. A new species of the genus Polyip- nus (Stomiformes, Sternoptychidae) from Su- ruga Bay, Japan.—Japanese Journal of Ichthy- ology 37(2):95—97. Baird, R. C. 1971. The systematics, distribution, and zoogeography of the marine hatchetfishes (Fam- ily Sternoptychidae).—Bulletin of the Museum PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of Comparative Zoology, Harvard University 142(1):1-128. ——.. 1986. Tribe Sternoptychini. Pp. 255—259 in M. M. Smith and P. C. Heemstra, eds., Smith’s sea fishes. MacMillan South Africa, Johannes- burg, 1047 pp. Borodulina, O. D. 1979. Composition of the ““Polyip- nus spinosus complex” (Sternoptychidae, Os- teichthyes) with a description of 3 new species of the group.—Voprosy Ikhtiologii 19:198—208. Gilbert, C. H. 1905. The deep-sea fishes of the Ha- waiian Islands.—United States Fish Commis- sion, Bulletin 23:575—713. Harold, A. S. 1989. A new species of Polyipnus (Sto- miiformes: Sternoptychidae) from the Coral Sea, with a revised key to the Polyipnus spi- nosus complex.—Copeia 1989:871—876. . 1990a. Polyipnus danae n. sp. (Stomiiformes: Sternoptychidae): a new hatchetfish species from the South China Sea.—Canadian Journal of Zoology 68:1112—1114. 1990b. Redescription of Polyipnus fraseri Fowler, 1934 (Stomiformes: Sternoptychidae), with remarks on paedomorphosis.—Proceed- ings of the Biological Society of Washington 103:509-515. . 1994. A taxonomic revision of the sterno- ptychid genus Polyipnus (Teleostei: Stomuifor- mes), with an analysis of phylogenetic relation- ships.—Bulletin of Marine Science 54:428— 534. . 1998. Areas of endemism and historical in- ference in biogeography. Pp. 148-155 in A. C. Pierrot-Bults, S. van der Spoel, B. J. Zahuranec, & R. K. Johnson, eds., Pelagic biogeography. UNESCO, Intergovernmental Oceanographic Commission, Workshop Report No. 142. , & S. H. Weitzman. 1996. Interrelationships of stomiiform fishes. Pp. 333-353 in M. L. J. Stiassny, L. R. Parenti, & G. D. Johnson, eds., The interrelationships of fishes. Academic Press, London, 496 pp. Last, P, & A. S. Harold. 1994. Polyipnus latirastrus, a new and unusual hatchetfish (Teleostei: Ster- noptychidae) from the western Pacific.—Copeia 1994:210—215. Leviton, A. E., R. H. Gibbs, Jr, H. Heal, & C. E. Dawson. 1985. Standards in ichthyology and herpetology: Part I. Standard symbolic codes for institutional resource collections in herpe- tology and ichthyology.—Copeia 1985:802— 832. Potthoff, T. 1984. Clearing and staining techniques. Pp. 35-37 in H. G. Moser, W. J. Richards, D. M. Cohen, M. P. Fahay, A. W. Kendall, Jr., & S. L. Richardson, eds., Ontogeny and systematics of fishes—American Society of Ichthyologists VOLUME 111, NUMBER 4 and Herpetologists, Special Publication Number 1:1—760. Schultz, L. P. 1938. Review of the fishes of the genera Polyipnus and Argyropelecus (Family Sterno- ptychidae), with descriptions of three new spe- cies.—Proceedings of the United States Nation- al Museum 86:135—155. . 1961. Revision of the marine silver hatchet- fishes (Family Sternoptychidae).—Proceedings of the United States National Museum 112:587— 649. . 1964. Family Sternoptychidae. Pp. 241—273 in H. B. Bigelow, C. M. Breder, D. M. Cohen, G. W. Mead, D. Merriman, Y. H. Olsen, W. C. Schroeder, L. PB. Schultz, & J. Tee-Van, eds., Fishes of the western North Atlantic, Part 4, Isospondyli—Sears Foundation for Marine Re- search, Memoir Number 1, 599 pp. Taylor, W. R., & G. C. Van Dyke. 1985. Revised pro- 953 cedures for staining and clearing small fishes and other vertebrates for bone and cartilage study.—Cybium 9:107—119. Weitzman, S. H. 1974. Osteology and evolutionary re- lationships of the Sternoptychidae, with a new classification of stomiatoid families.—Bulletin of the American Museum of Natural History 53: 327-478. Wessel, J. H., III. 1997. The Myctophidae of the So- mali Current. Unpublished MSc. thesis, Univer- sity of Charleston, South Carolina, two vol- umes, 374 and 250 pp. , & R. K. Johnson. 1998. The Sternoptychidae of the Somali Current region of the western In- dian Ocean: an introduction to Somali Current mesopelagic fish studies. Pp. 372-379 in A. C. Pierrot-Bults, S. van der Spoel, B. J. Zahuranec, & R. K. Johnson, eds., Pelagic biogeography. UNESCO, Intergovernmental Oceanographic Commission, Workshop Report No. 142. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):954—985. 1998. A review of western north Atlantic species of Bembrops, with descriptions of three new species, and additional comments on two eastern Atlantic species (Pisces: Percophidae) Bruce A. Thompson and Royal D. Suttkus (BAT) Coastal Fisheries Institute, Louisiana State University, Baton Rouge, Louisiana 70803-7503; (RDS) Tulane University Museum of Natural History, Belle Chasse, Louisiana 70037 Abstract.—The Atlantic members of the genus Bembrops were examined from approximately 700 specimens. Meristic, morphometric, and pigment char- acters were used to redescribe six species, B. anatirostris, B. gobioides, B. greyi, B. heterurus, B. macromma, and B. magnisquamis. Three species, Bem- brops ocellatus, B. quadrisella, and B. raneyi are described as new. New in- formation on sexual dimorphism, the cephalic canal system and vertebral num- . ber is presented. Distribution maps for all nine species are shown. Bembrops ocellatus and B. quadrisella are widespread in the Caribbean Sea. Bembrops raneyi is known only from the Bahama Islands. The three new species are illustrated. The American members of the genus Bembrops were reviewed by Ginsburg (1955); three of the four species he de- scribed as new; in addition, he reviewed the literature and characterized the family Per- cophidae (=Percophididae). The family was also characterized by Grey (1959) and detailed descriptions of B. anatirostris, B. gobioides, and B. macromma were present- ed, along with figures of B. anatirostris and B. macromma. More recently, Das & Nelson (1996) pre- sented a world-wide revision of the perco- phid genus Bembrops, recognizing a new species from the eastern Atlantic. They re- viewed the eighteen nominal species in the genus, relegating five into synonymy. A wealth of unstudied material of the various species is available from Cornell University (CU); Tulane University, Muse- um of Natural History (TU); University of Florida (UF); and United States National Museum (USNM) and we present addition- al data on the four species known to Gins- burg, plus data on two eastern Atlantic spe- cies, and describe three new species from the western north Atlantic. During this study we have become aware that much remains to be done in systemat- ics, life history, sexual dimorphism, and distribution of the species within the genus Bembrops. We intend to pursue the study of Atlantic populations and expand our study to include related Indian and Pacific Ocean species. However, because of practical con- siderations the main purpose of this paper is to provide names for the three new forms. We follow Ginsburg’s (1955) placement of Bembrops in the family Percophidae (=Percophididae). Future research may prove the necessity of elevating the subfam- ily Bembropinae (Nelson 1976, 1984, 1994) to family status. The new species de- scribed herein fit the characters for the fam- ily and genus as given by Ginsburg (1955), Nelson (1976, 1978, 1984, 1994) and Das & Nelson (1996). Methods and Materials Ginsburg (1955) pointed out the difficul- ties involved in studying the species of VOLUME 111, NUMBER 4 Bembrops because most are captured in bottom trawls and often specimens suffer broken fin rays and loss of some or most of their body scales. Difficulties in measuring and making some counts are increased when specimens have twisted, contorted bodies, and widely flared branchiocranial structures. Some of our lateral-line scale counts are estimates based on counts of scale “‘pockets’’ and some of the variation presented in the morphometrics is due to the use of distorted specimens. We follow Ginsburg (1955) and Hubbs & Lagler (1958, 1964) in fin ray and gill raker counts. We counted rudimentary ele- ments on both upper and lower limbs of the first left gill arch, and the raker at the angle of the arch was included in the count of the lower limb. Although Das & Nelson (1996) cited Hubbs & Lagler (1964) in their Meth- ods and Materials section, our gill raker and fin ray counts do not always agree. Depth of body was measured at the ori- gin of first dorsal fin. Caudal peduncle was measured at its least depth. The head was measured from the anterior point of the rounded lobe near the end of the snout rath- er than from the indented midpoint to the posterior end of the extended fleshy oper- cle. The snout was measured from this Same point to the anterior rim of the orbit. The eye (orbit) was measured horizontally between the most anterior and posterior fleshy rims of the orbit. The interorbital measurement is the least bony interorbital distance. Upper jaw length was measured from its anterior tip to the bony posterior tip of maxilla (not including tentacle). The lower jaw measurement was interpreted as the distance between anterior tip of dentary to the posterior bony tip of maxilla (not in- cluding tentacle). Left and right tentacles are seldom of equal length and so we measured both whenever feasible, especially for species of which we had few specimens. The tentacles are quite elastic and before measurement each tentacle was gently stretched straight 955, with a fine forceps, allowed to contract, then measured. Pigmentation patterns in the first dorsal fin were determined for the nine Atlantic species of Bembrops included in our study (Fig. 1). Pigmentation on other fins and on body is also described. Vertebral counts were made from radio- graphs of 333 specimens taken primarily by the Radiology Department, Louisiana State University School of Veterinary Medicine. Counts were made on 113 Bembrops an- atirostris, 178 B. gobioides, 60 B. greyi, 75 B. heterurus, 67 B. macromma, 8 B. mag- nisquamis, 39 B. ocellatus, 123 B. quadri- sella, and 29 B. raneyi. Measurements were made on 50 B. an- atirostris, 50 B. gobioides, 62 B. greyi, 50 B. heterurus, 45 B. macromma, 6 B. mag- nisquamis, 14 B. ocellatus, 16 B. quadri- sella, and 19 B. raneyi. Proportional mea- surements were made with a needle-point dial calipers and recorded to the nearest 0.1 mm. Meristic data are presented in Tables 1-4 and morphometrics are presented in Tables 5-10. In Tables 5-7 and 10 the proportional measurements are expressed in thousandths of standard length (SL). Table 8 shows fre- quency distributions of ratios of head width divided by head depth, Table 9 contains fre- quency distributions of snout length divided by orbit length. Table 11 presents a sum- mary of typical fin pigmentation patterns for the nine Atlantic species with sexual di- morphism noted. We investigated the full extent of the ce- phalic portion of the lateral-line sensory system, which was only briefly mentioned by Ginsburg (1955) and Grey (1959). The supracleithrum and posttemporal bone were dissected from the right side of five species (B. anatirostris, B. gobioides, B. greyi, B. heterurus, and B. macromma) and mea- sured, and expressed as percent of SL. The connection of the cephalic canal system to the lateral-line was examined to determined to what degree the canal is enclosed by bone. 956 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Spinous dorsal fin pigment pattern in nine Atlantic Bembrops: A) B. anatirostris, B) B. greyi, C) B. gobioides, D) B. heterurus, E) B. macromma, F) B. magnisquamis, G) B. ocellatus, H) B. quadrisella, and I) B. raneyli. The alimentary tract was dissected on 3— Collection depths were obtained from 4 specimens of B. anatirostris, B. gobioi- cruise reports and reported in meters (m). des, B. greyi, B. heterurus, and B. macrom- Maps in Williams (1968) were used for lo- ma and pyloric caeca measured; length of cating collecting sites of the two eastern At- caeca is given as percent of SL. lantic Bembrops. Table 1.—Frequency distribution of the number of dorsal, anal, and pectoral rays in nine Atlantic species of Bembrops. Dorsal rays Anal rays Pectoral rays Species 14. 15 16 i7uis io” 1 mig.) 10. 22 °03) 24. 25) (260 io OCONEE B. anatirostris 8 105 20 Oy Il BG 59 31 4 B. gobioides 9 163 6 4 163 9 30, dh e33 selene B. greyi 10 48 2) 26 34 4 44 12) B. heterurus 6 67 2 1 69 5 25 50 B. macromma 66 16 50. 61 4 45 18 B. magnisquamis 7 1 8 PRB 64 B. ocellatus 8 28 2 7 DOD, 4 23 11 1 B. quadrisella 17 101 5 37 82 8 53 54 8 B. raneyi 3 25 1 9 20 1 3 18 6 | VOLUME 111, Table 2.—Frequency distribution of the number of lateral-line scales in nine Atlantic species of Bembrops. Scales NUMBER 4 70 69 Species B. anatirostris B. gobioides B. greyi 11 15 19 18 13 B. heterurus 10 B. macromma 1 4 B. magnisquamis B. ocellatus 12 10 B. quadrisella B. raneyi 957 Species accounts and distribution maps are based almost entirely on our own ob- servations, not compilations of previous published information, so nearly all speci- mens used in this study have never been published upon. Exceptions are a few spec- imens from the National Museum of Nat- ural History (USNM) and University of Florida (UF) cited in Das & Nelson (1996). Several Pacific Bembrops were examined for comparison to the nine Atlantic species: B. curvatura CAS 32808, 32976, 33100, 34393, 34589, and 88698 (25); B. filifera CAS 89509 (8); B. nematopterus USNM 347251 (1); B. platyrhynchus CAS 88685, USNM 345189 (5); B. sp. nov. CAS 88680, USNM 345190 (7). We recommend the use of “‘duckbill’” for the common name of members of the genus Bembrops to avoid confusion with the commonly used name “flathead” as applied to the family Platy- cephalidae (Eschmeyer 1990). We suggest common names for the nine Atlantic spe- cies. Genus Bembrops Steindachner, 1876 We examined the cephalic sensory pore system in detail in Bembrops anatirostris, B. ocellatus, and B. raneyi and found the general configuration of the canals to be nearly identical. We found that canals and pores were more discernible on those spec- imens that had lost patches of scales and pigmented skin from the head. Figure 2A is a composite sketch representing several specimens. The most anterior pore is on either side near anterior tip of snout and the canals ex- tending posteriorly from these two pores gradually converge toward the anterior por- tion of interorbital area. These two canals (left and right) pass medial to the anterior and posterior nares, respectively, and the second set of pores project laterally at this place and may easily be confused with the narial openings. The third set of pores is in a staggered arrangement, with respect to each other, at the anterior interorbital area 958 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 3.—Frequency distribution of the number of gill rakers in nine Atlantic species of Bembrops. Upper limb Lower limb Total, both limbs Species 3 4 5) 6 12 ts} 14 15 16 17 «+18 15 16 17 18 19 20 21 22: 93 B. anatirostris 12, 97 A PDD, X39). KS) AL We 73 Mk) 1 B. gobioides Wy NS 3} 1 49 107 18 1 1 See SS. Ie 2 B. greyi 6 49 5 Ay ily 3) 1 6 14 36 3 B. heterurus i S7/ BUS) A'S) 3 1 10 24 37 2 B. macromma Dm DAs 3) VADs 6 1 4 psa) = 35)» 2 B. magnisquamis 1 7 Dinan | Q'S 2) nt et ES ee B. ocellatus 1 33% | 12 18 5 1 ie SiS: 1 B. quadrisella 106 See 27 OT 18 41 27 197 i B. raneyi Dim DAS 6 18 5) DB a WS 8 where the two canals (left and right) are separate but adjacent to each other. Between the anterior and posterior interorbital areas the two canals seem to join to form a single canal and at the posterior interorbital area there is a centrally located single pore. We do not know that this arrangement is typi- cal, however an air jet placed at the single pore forced liquid out of both pores at the anterior interorbital area and the pores near the narial openings. Immediately posterior to the single, in- terorbital pore the canal divides into a left and right branch and respectively circles around posterior rim of orbit. About half way around the posterior rim of orbit there is a branch canal that courses posteriorly and at the occipital area a short side branch projects medially, but right and left side branches do not join across the occiput. Af- ter the small side branch, the main (branch) canal continues to extend posteriorly and Table 4.—Frequency distribution of the number of vertebrae in nine Atlantic species of Bembrops. Species 2 OOO) N x SD B. anatirostris 1 41 42 27.98 0.15 B. gobioides 2 33) a 2M O73) B. greyi 8 46 54 29.85 0.36 B. heterurus 49 1 50 28.02 0.14 B. macromma 3 Sy) G2 27953 022 B. magnisquamis 8 8 29.00 — B. ocellatus 2 12 14 29.87 0.34 B. quadrisella © Sil 337 A753) (0),337/ B. raneyi 4 25 29 29.86 0.35 connects with the anterior end of each lat- eral-line canal. The two systems meet or join under the tip of a diagonally directed spine (posttemporal of some authors) and the anterior keeled lateral-line scale. The number of pores in the infraorbital canal posterior to the orbit varies in several spe- cies, having either 3 or 4 pores. There is also a mucous canal and pore system associated with the preoperculo- mandibular area. This system is not con- nected with the cephalic system described above. The preoperculomandibular system extends from upper end of preopercle, ven- trally and then anteriorly to anterior tip of lower jaw. The number of pores seem to be variable. Figure 2B illustrates the right preoperculomandibular canal of a B. ocel- latus with nine pores. Thus we find the ce- phalic lateral-line system of Bembrops is somewhat abbreviated in comparison to that of the genus Acanthaphritis, subfamily Hemerocoetinae (Suzuki & Nakabo 1996) with Bembrops having the infraorbital canal very short, extending less than halfway to the anterior margin of the orbit and the preoperculomandibular branch not connect- ing with the lateral-line canal (Fig. 2A, B). The cephalic canal system that passes through the posttemporal bone (ptt) and along the top of the supracleithrum (scl) in connecting with the lateral-line varies among the five Atlantic species examined. The size of each bone, expressed as percent of SL, and the degree to which the canal is 959 VOLUME 111, NUMBER 4 : ; Bi van- losed by bone differs as follows: B Aes 1H ew €ncios 1 enclosed, longest ana ttY WwMmmM Oo atiros Fi A 3A); B. All aim Gon amoa tt and scl, 6.5% and 5.7% ( e hortest N = Pp ? roove, sho joides—canal an open g ‘ ‘ gobioides: d 4.2% (Fig. 3B); B. 5 = 2S eS ain OB Se 2 wc ptt and scl, 5.3% an romma are So MEMES mt greyi, B. heterurus, and B. leeeeokn 3 Hl Pe ‘ | more or less oS & lar with cana ve Rf 26 +t very sim ith a single pore along : ASy+Soa losed with Fea} tom S ss a i ees 32 a pletely enc ; atl att aul scl, 6.8% ee a Ase le ah: ay ak intermediate-sized p : a Ed S length, inte 5% 4 5 SrnaArr avons engtn, 2%, 6.5% and 5. 0, 5} Bs ai eS ”n a Oo Sy oo ee and 5 3%, 6.4% and 5. 5 h ie a single S tS ae < —E). There i 5 respectively (Eig. 3C eee posteriorly o ORR EOS AR RES SS large, exposed spine Sealine Aeeal : 8 gedaqurongnay fio teeat erie 8 of the cephalic canal. Q. 2 on =Oot oO EMR a} Se os ee = Bembrops anatirostris mei = | gn Longnose duckbi < 5 ) S SONTADS 4 VI, 14 E ; xf RAS Be 2 a aas 7 a Dorsal fin in 113 specimens has Anal 2 es Ted Ya ae at Il with 15 rays. An a | aN Ont 4 nt Oo ft rays, nearly a 5 RA Kohan eee Che reeeenere ys with a strong mode of 3 Ss fin has 17 or ne to 29 2 RS 18. Pectoral fin rays range me UES 3 Q HAGKSSTEH aes with the mode at 27 ila ai ay 5 Bh pases Lateral-line scales See ange from om ie ill rakers r. Le Table 2). Gill > X = 63.3 ( , 16 on lower a 1s In daegenesgnoe Tee te ie ee eran fe ey es les 2 eae Mes d total coun A g limb an almost E e (Table 3). Number of nace st ae ea < Axe ; ount oO 5 : eee sce always 28, only a single c Ce Bill seas eesial Adsdaannads 4 a 5 f b ngest 2 “| 2a cl Gre BBS S = Bembrops anatirostris has : pe Ne all : < nout of the nine Atlantic eee ae wis eas S 2 1 the sn z HRHODAAWONOM + S imens examined having bles I) =SHeGatecsa 5 ON spec bit length (Table SWIG Phe ol coe tA os than the orbi : s Bl Gad 6 x length greater long thin & +4 =I 5 9) Bembrops anatirostris has a lo & t z eae tacle, attenuate a ) . xillary ten ; 2 |< aon arte ERE oe enx ae f tentacle of the nine o Soi | & = See aS . he longest 2 lee eat e Sea Meee eet ae oe Aa ee LAO) * = G onsidere in 2 i © |:. species C hows a rudimentary ten Solis 2 NL DeRES Richards (1990) s is and a 5 Nowe AHRSOHOSHSH SL B. anatirostris a E es Sn ein tania Ke tacle on a 9.4 mm ; mm SL (al- 2 5 aD OAL MoAO Ow ow d tentacle on a 15 2 [a7 = o+ al lon om S sa) well-forme ; ipilit these figures 5 OF h there is a possibility h = vis B eyi, also known from the =| be . ran ) SS ie] may é | ro : orida). < s fa 2s Siaisof st anatirostris is the only one of o 3 ° = bo Bb By Bembrop ies in this study that has any © ie 3 = 55 5 ine species in t 1 a E SI 2 al a OS = the nine sp ines: the second dorsa | 2 Si hese haees longed dorsal spines; e Be “| BEPRSS Peas euCnOn e males. We can ® . pugosresas 5S spine is prolonged in larg 1996) report- 2 Ss S54 0 = 2 fo GRE Es 2 2 Dea ee ea arene & AgGeseEaSaSS YN 960 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 6.—Proportional measurements (expressed in thousandths of standard length) for two eastern Atlantic species of Bembrops. B. greyi n = 62 Proportion Range x SL mm 54.9-223.4 125.6 Body depth 96—156 118 Caudal peduncle depth 48-63 55 Head length 366-428 388 Head depth 96-137 114 Head width 138-172 153 Interorbital width 5—10 7 Snout length 82-122 105 Orbit length 100-125 112 Upper jaw length 138-175 155 Lower jaw length 152-191 171 Postorbital length 156-190 174 ing elongation of the third spine of first dor- sal of this species since all males examined in this study had only the second spine elongated. There is a moderate correlation between length of specimen and length of spine. Twenty-one males that range from 147 to 230 mm in standard length have sec- ond dorsal spines (expressed in thousandths of standard length) that range from 172 to 381, X = 265. The elongate spine possesses a narrow “‘flag’’ of black epidermis along the posterior margin from the margin of the fin to the tip of the spine. The first two membranes of the spinous dorsal fin of B. anatirostris are a dull black, the remaining membranes are typically clear (Fig. 1A). Small specimens have an upper basal caudal spot but neither adult males nor females retain a distinct caudal spot. Bembrops anatirostris has three long py- lone caecayal225 91322 vandal S(leit to right) % of SL. The caeca extend past the posterior end of the undistended stomach. Distribution/Depth.—Northern Gulf of Mexico; Gulf of Campeche; also from northwestern edge of Little Bahamas Bank, western Caribbean Sea off Rosalind Bank and southern Nicaragua; southwestern Ca- ribbean Sea off Panama; western end of Puerto Rico and Atlantic Ocean east of Dominica. Off South America—Colombia, B. heterurus n=5 SD Range xX SD 42.27 80.7—214.5 145.8 35.03 13.14 121-167 137 11.07 3.32 51-66 57 3.17 13.59 351-398 377 10.07 8.60 109-156 124 9.78 7.36 133-191 162 12.82 1.18 7-12 10 1.18 10.35 106-124 114 4.05 6.75 74-103 93 5.88 10.86 140-160 152 5.47 10.79 158-187 7/3} 6.85 8.20 148-178 167 5.69 eastern Venezuela, Guyana, and Suriname (Fig. 4). Additional records in Ginsburg (1955), Grey (1959), Uyeno et al. (1983), Boschung (1992), and Das & Nelson (1996). Bembrops anatirostris is the shal- lowest occurring species of the genus in the Atlantic. The depth range of our B. anati- rostris is 82-549 m, with most specimens from 320 m or less. Ginsburg (1955) re- ported the species from 110—366 m, while Grey (1959) listed her material from 139-— 366 m, with most specimens from 220 m or less. Bullis & Struhsaker (1970) reported the highest density of B. anatirostris from 185—274 m, with few specimens from deep- er waters. Das & Nelson (1996) listed the species from 100 to over 350 m. We ex- amined one collection of B. anatirostris taken with B. magnisquamis off Costa Rica and one collection taken with B. quadrisella off Venezuela. Material examined.—TU 26887 (1): Oregon 156; 27°22'N, 96°08’W; 180 m; 27 Nov 1950. TU 2732 (10): Oregon 278; 29°49'N, 85°45'W; 205 m; 24 Feb 1950. TU 10689 (7): Anna Inez 644; 28°52’N, 88°59'W; no depth recorded; 7 Jul 1955 [pyloric caeca dissection]. TU 10995 (A): Oregon 1094; 27°10'N, 96°20'W; 274 m; 5 Jun 1954. TU 12891 (4): Oregon 1095; 27°10'N, 96°17'W; 320 m; 5 Jun 1954. TU 12896 (2): Oregon 1093; 27°03'N, VOLUME 111, NUMBER 4 961 . ABAAM OOH TEND 96°16’W; 384 m; 4 Jun 1954. TU 12962 2, OSM ACH ASK AHH (1): Oregon 895; 28°47’N, 85°19'W; 117 m; 7 Mar 1953. TU 12981 (2): Oregon 864; a 29°19'N, 86°04’W; 150 m; 31 Oct 1953. TU SPA aHSOnMSesege 17609 (3): Oregon 2000; 07°55’N, an 57°30'W; 82 m; 5 Nov 1957. TU 90207 (1): Bul [oo Oregon-II 13167; 29°15.7'N, 88°09'W; 128 slg) S8nRSR-S2S68| mm: 3 May 1973. TU 90224 (1): 29°14.7'N, Z)—oeersacgezadg| 88°08.5'W; 165 m; 3 May 1973. TU 90241 © ae east (8): Oregon-II 13170; 29°14'N, 88°09.3/W: al 183 m; 3 May 1973. TU 180139 (7): 3 Swurowrartarng Oregon 1985; 09°41’N, 59°47'W; 274 m; 3 a =] Sirs Pan ra aa Nov 1957. TU 180140 (5): Oregon 2012; & 07°34'N, 54°19’W; 274 m; 8 Nov 1957. TU 5 DANTRT HARKS 180141 (4): Oregon 2203; 29°13.5'N, “ Se GA aE ks 88°12'W; 229 m; 26 Jun 1958. TU 180142 a (1): Silver Bay 441; 27°39'N, 79°15'W; s CRS APSR ees ah hk 503-549 m; 9 Jun 1958. TU 180337 (3): m ond n te -—” HON Oo Oo ° ’ ° ’ a (ES Sarah ASiay Clay eS CNTs Oregon 1984; 09°45'N, 59°45’W; 366 m; 3 2 Be Nov 1957. TU 180338 (8): Oregon 1989; B | SU Itc eum ele Se wes 09°45’N, 59°45’W; 366 m; 4 Nov 1957. TU Ste [B)G546555aS°255) 180339 (10): Oregon 2005; 07°37'N, < 2) CRSSRACSRAAR! 54°50’W; 366 m; 6 Nov 1957. TU 180340 = 3 (1): Oregon 2023; 07°15’N, 53°21'W; 247 3s , m; 9 Nov 1957. TU 180341 (1): Silver Bay E g rs baonagonnoagy 175; 28°0S'N, 90°52'W; 183 m; 20 Sep z Re ae ge Both dear 1957. TU 180342 (5): Oregon 2022: 2 07°15'N, 53°25’W; 210 m; 9 Nov 1957. TU 2 p| BSRSIALRASSZ) 180343 (8): Oregon 1983; 09°53’N, 3 = = 59°53'W; 229 m; 3 Nov 1957. TU 180344 2 (2): Oregon 1981; 10°03'N, 60°01'W; 366 2 «le 9ege5neggueg] m3 Nov 1957. TU 180345 (2): Silver Bay Sovalkes Sh BSE ah aig 154; 28°58'N, 84°44’W; 139-148 m; 22 3 [ss Aug 1957. TU 180346 (1): Oregon 1884; Oe Als S seh hho eaeer 16°53’N, 81°22’W; 411 m; 23 Aug 1957. CMe i tnt a TU 180347 (1): Silver Bay 100; 29°10’N, a4 OO FANADANHNADAA OHH g EDT OUD NOMI 85°48’W; 101-130 m; 26 Jul 1957. TU 180348 (1): Oregon 1986; 09°39’N, : 21 on 59°47'W; 183 m; 4 Nov 1957. USNM g e] BABS SRALH! 304921 (2): Oregon 4838; 11°09’N, = 74°24'W; 329 m; 16 May 1965. USNM 6 2 343811 (2): Oregon 2658; 18°26’N, 5 3 Z eam 67°11.5'W; 320 m; 7 Oct 1959. Examined £ g 3 3s et 2 Sp for distribution information (Fig. 4): USNM ! S| gaaae 5 creas 3 231943 (2), 278482 (4), 308117 (4), 2 | ereeeis es s)8 502) || 347203' (6), 347222(1), 347274), iS Bole = aisle )8 2 8 347275 (2), 347276 (2), 347277 (3), AgoOHEEEGODSES BAT DAISY (i); 6 347279. (Os 847280116), 962 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 8.—Frequency distribution of head width divided by head depth in nine Atlantic species of Bembrops. Standard length Species in mm n 1.1 B. anatirostris 65.8-246.0 50 B. gobioides 84.6—217.4 50 B. greyi 54.9-223.4 62 1 B. heterurus 80.7—214.5 50 1 B. macromma 77.3-190.0 45 B. magnisquamis 72.0-104.2 6 B. ocellatus 90.9-190.0 14 B. quadrisella IDES 235-5 16 B. raneyi 69.1—167.3 19 347281 (4), 347282 (4), 347283 (3), 347284 (2), 347285 (8), 347286 (1). Bembrops gobioides (Goode, 1880) Goby duckbill Dorsal fin in 178 specimens has VI, 16— 18 soft rays, with a strong mode of 17 rays. Anal fin rays vary in number from 17 to 19, mostly 18. Pectoral fin rays range from 26 to 30, with a strong mode of 27 (Table 1). Lateral-line scales range from 60 to 70, X = 64.6 (Table 2). Gill rakers range from 4 to 6 on upper limb, 13 to 17 on lower limb, and total counts range from 17 to 22, usually 19 or 20 (Table 3). Number of ver- tebrae is almost always 30 (35 of 37), two counts of 29 (Table 4). Based on 50 specimens ranging from 84.6 to 217.4 mm in SL, five have the orbit length equal to the length of snout but most specimens have the snout longer than the eye (Tables 5, 9). An additional small spec- Ratio 1.2 1.3 1.4 1.5 1.6 ea 9 20 18 3 6 25 15 4 W 32 17 3 2 9 BY 12 1 2 29 14 1 — 1 3 1 1 9 3 1 1 4 7 4 4 11 3 1 imen (84 mm SL) not included in Tables 5 and 9 has the orbit slightly greater in length than length of snout. This observation matches that presented by Ginsburg (1955, Table 4) for an 81 mm SL specimen. Bembrops gobioides typically has 17 soft dorsal rays, 18 anal rays, 27 or 28 pectoral rays, more than 60 lateral-line scales, and 30 vertebrae all among the highest values for these characters for the nine Atlantic species. Its 30 vertebrae contrasts with B. anatirostris which typically has 28 verte- brae. The caudal peduncle of B. gobioides is less deep than that in B. anatirostris. There is a more pronounced change in rel- ative size of orbit and snout with respect to size of specimen than was observed for B. anatirostris. No small specimens of B. an- atirostris has the orbit exceeding or equal to length of snout, whereas B. gobioides specimens between 80 to 85 mm in SL have the orbit equal to or greater than length of Table 9.—Frequency distribution of snout length divided by orbit length in nine Atlantic species of Bembrops. Standard length Species in mm n 0.7 0.8 . anatirostris 65.8—246.0 50 gobioides 84.6-217.4 50 greyi 54.9-223.4 62 5) 11 . heterurus 80.7—214.5 50 macromma . Magnisquamis . ocellatus . quadrisella . raneyi 72.0-—104.2 6 90.9—190.0 33) 127.5—235.3 94 Db bbb bb & & 77.3-190.0 45 1 10 69.1—167.3 19 3 Ratio 0.9 1.0 1.1 U2 1.3 1.4 1.5 1.6 2 5 14 18 10 1 5 17 21 7 15 19 12 24 iL) 4 1 VOLUME 111, NUMBER 4 Table 10.—Maxillary tentacle length (expressed in thousandths of standard length) in nine Atlantic spe- cies of Bembrops. Species n Range X SD B. anatirostris 53 33-71 51 9.42 B. gobioides 48 20-37 28 4.20 B. greyi 121 15-51 33 7.78 B. heterurus 98 12-50 30 8.83 B. macromma 87 7-27 18 4.60 B. magnisquamis 12 17-49 37 9.70 B. ocellatus 27 19-39 26 5.33 B. quadrisella 30 14-27 21 3.50 B. raneyi 38 31-59 41 7.47 snout. The maxillary tentacle of B. gobioi- des is small to moderate in length and is triangular in shape, whereas B. anatirostris has a very long tentacle that has an atten- uate distal portion. The spinous dorsal fin pigmentation of B. gobioides is similar in some respects to that described for B. anatirostris; the first two membranes are blackened, however there are additional wedges of black pigmenta- tion at the distal ends of spines 3—6 (Fig. te): Bembrops gobiodes has three short py- loric caeca, 6.0, 5.4, and 6.2 (left to night) % of SL. Distribution/Depth.—Northwestern, northcentral and northeastern Gulf of Mex- ico; Dry Tortugas; southern Gulf of Mexi- co, north of Yucatan; off east coast of Flor- ida; western Little Bahamas Bank; no re- cords from Caribbean Sea (Fig. 5). Addi- tional records in Ginsburg (1955), Grey (1959), Boschung (1992), and Das & Nel- son (1996). The depth range of our Bem- brops gobiodes is 329-549 m, with most specimens taken between 350 to 450 m. Ginsburg (1955) listed this species from 119-512 m and Grey (1959) reported it be- tween 338 and 438 m. Das & Nelson (1996) reported it from about 100 to over 700 m. Bullis & Struhsaker (1970) reported the highest densities of B. gobiodes from the western Caribbean in two depth ranges, 276—366 m and 459-549 m, but these prob- ably represent a composite of two of our 963 new species since we have no valid records of this species from the Caribbean Sea. Material examined.-TU 1709 (6): Oregon 61; 29°04'N, 88°30’W; 357 m; 2 Aug 1950. TU 2692 (1): Oregon 62; 29°01'N, 88°30’; 424 m; 2 Aug 1950. TU 2729 (4): Oregon 162; 27°18'N, 96°09'W; 366 m; 28 Nov 1950. TU 2730 (5): Oregon 163; 27°15'N, 96°00’W; 430 m; 28 Nov 1950. TU 11714 (8): Oregon 1520-80; 29°10'N, 88°10’ W; 366—457 m; 16-19 May 1956. TU 12869 (4): Oregon 1107; 29°03'N, 88°25'W; 384-430 m; 15 Jun 1954 UP A2s8oie ©). Orecony 093. 27°03'N, 96°16'W; 384 m; 4 Jun 1954. TU 12943 (13): Oregon 1091; 26°41'N, 96°20'W; 366-384 m; 3 Jun 1954. TU 14779 (2): Pelican 60; 28°29'N, 79°54’W; 293-347 m; 11 Jun 1956. TU 14810 (1): Pelican 66; 28°33'N, 79°52'W; 357-371 m; 13 Jun 1956. TU 17073 (25): Combat 332; 28°55'N, 79°56'W; 329 m; 31 May 1957 [pyloric caeca dissection]. TU 35788 (3): Oregon 4367; 24°31’N, 83°30'W; 375 m; 5 Aug 1963. TU 35802 (3): Oregon 4367; 24°31'N, 83°30'W; 375 m; 5 Aug 1963. TU 35803 (5): Oregon 4368; 24°31'N, 83°30'W; 375 m; 5 Aug 1963. TU 35805 (5): Oregon 4368; 24°28'N, 83°26'W; 384 m; 5 Aug 1963. TU 35832 (3): Oregon 4371; 24°28'’N, 83°27’W; 375 m; 6 Aug 1963. TU 35863 (2): Oregon 4372; 24°29'N, 83°32'W; 375 m; 6 Aug 1963. TU 90280 (1): Oregon-II 13208; 29°16.1'N, 87°38.4'W; 457 m; 4 May 1973. TU 90300 (2): Oregon-II 13210; 29°15.6'N, 87°45.0'W; 411 m; 4 May 1973. TU 90320 (1): Oregon-II 13218; 29°14.5'N, 87°44’W; 494 m; 5 May 1973. TU 90342 (6): Oregon-II 13222; 29°10.4’N, 88°07.9'W; 366 m; 5 May 1973. TU 90398 (7): Oregon-II 13254; 28°54.7'N, 88°45.6'W; 347 m; 7 May 1973. TU 90406 (18): Oregon-II 13256; 29°00'N, 88°35.5'W; 347 m; 7 May 1973. TU 90425 (10): Oregon-II 13260; 28°55.2'N, 88°40'’W; 411 m; 7 May 1973. TU 90468 (5): Oregon-II 13332; 24°23.4'N, 83°24.1’W; 402 m; 21 May 1973. 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OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. B. Right preoperculomandibular canal of B. ocellatus. Ptts = posttemporal spine, L1 = lateral-line. 83°09.9'W; 402 m; 22 May 1973. TU 90503 (13): Oregon-II 13344; 24°16'N, S2e5 22 Wa AOD anise 2) (May 197Saat 90565 (1): Oregon-II 13362; 24°09'N, 82°31'W; 549 m; 23 May 1973. TU 180143 Cs Silver Baye221 29221 Ne SO,0Gnv- 329-347 m; 22 Nov 1957. TU 180144 (1): Silver Bay 212; 29°59’N, 80°07'W; 366 m; 20 Nov 1957. TU 180145 (1): Silver Bay DIA 2929 N.S SO%09 We 329) me 245 Nov 19572 TU, is0l4e, (@): Silver Bay 217: 29°41'N, 80°08’W; 329-366 m; 21 Nov 1957. EW Ws0l47 “@)= Combat, 464: 27°51'N, 79°50'W; 393 m; 29 Jul 1957. TU 180148 (1): Combat 482; 29°28'N, 80°08'W; 347 m; 17 Aug 1957. TU 180149 (5): Combat 317; 29°07'N, 80°04’W; 366 m; 27 Apr 1957. TU 180150 (6): Combat 462; 27°14'N, 79°50'W; 384 m; 29 Jul Ie WU IKONS (ye Coiba S30 29°08'N, 80°03'W; 347 m; 31 May 1957. TU 180152 (4): Combat 314; 29°38'N, 80°11'W; 329 m; 27 Apr 1957. TU 180153 Cephalic sensory pore system: A. Composite of Bembrops anatirostris, B. ocellatus, and B. raneyi, (5): Combat 467; 28°36'’N, 79°54'W; 402 m; 30 Jul 1957. TU 180154 (2): Combat 479; 29°22'N, 80°06’W; 366 m; 17 Aug 1957. TU 180155 (1): Gombatwsie: 29°20'N, 80°04'W; 393 m; 27 Apr 1957. TU. 180156 ©@): Combat 463; 2 7-22.55 78°50'W; 402 m; 29 Jul 1957. TU 180349 (1): Combat 501; 29°36'N, 80°07’W; 384 Mla ep eos i Bembrops greyi Poll, 1959 Blackfin duckbill Dorsal fin in 60 specimens has VI, 15 to 17 soft rays, with a strong mode of 16 rays. Anal fin has 17 or 18 rays, with a greater frequency of 18 rays. Pectoral fin rays range from 25 to 27 in number, with a mode of 26 (Table 1). Lateral-line scales range from 45 to 52, X = 49.2 (Table 2). Gill rakers range from 3 to 5 on upper limb, 12 to 14 on lower limb, and total counts range from 15 to 19, usually 17 or 18 (Table 3). VOLUME 111, NUMBER 4 967 Fig. 3. Posttemporal and supracleithrum of five species of Bembrops: A) B. anatirostris, B) B. gobioides, C) B. greyi, D) B. heterurus, and E) B. macromma. Ptt = posttemporal, Scl = supracleithrum. ‘A PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 968 = 1 1 -4----}-+5 A SE --+-4-4--|-- av | | |. ISPANIOLA. BBE RI | 1 + Shoe oie a bad = 4 = + S. AMERICA -4----+-+-4- oe ae 5 ik: Distribution of Bembrops anatirostris (@), B. magnisquamis (&), and (*) both species. Fig. 4. ! a 2 f I ale: IL A : | tal “aa | ] — ——+ T ks r om : at! a | = <4} T | 1 — aft | | ! t el Jeol ae eee iS ae a ee ee ie if +—_1—+ “fe | ! iM +} aT I 4a . a2 | 22) 2 ee = | | = I _ i | fay ee Le Sy HL ns ! ne 3 =I < i G 4 a I *] i é ix : T a 1 hat B= Hy 1 1 Fiala Al 4--p-+-4-- = 1 | a + | 5 | | | a aa 3 Th 2 \ § ee =F Hake | 1 ae ; ‘q. +t “a > ail, ! | ae | v ih a 7 | : = pW S| He Se { ry § = ae = i j | | | a a 5 ot a 5 | pty oy —— a : | | | Panes | bese | mek ces He i ; Me H ! | } SS ee ee fa ee ele i < | H | a jess ote = 5 | ae ee Na i 1 T S : jt i Distribution of Bembrops gobioides (®), B. macromma (A). RIS: 2 VOLUME 111, NUMBER 4 Number of vertebrae is typically 30, how- ever eight have 29 (Table 4). Based on 62 specimens ranging from 54.9 to 223.4 mm SL; half (31 specimens) have the orbit length greater than length of snout; the orbit and snout lengths are equal in 19 specimens; and in 12 specimens the snout length is greater than the orbit length (Table 9). Based on 121 maxillary tentacle mea- surements (representing 62 specimens) the tentacle length in B. greyi is slightly greater than average among the nine species (Table 10), although B. anatirostris, B. raneyi, and B. magnisquamis have longer tentacles. In summary, Bembrops greyi has a low number of lateral-line scales with only B. magnisquamis having as low a lateral-line scale count. Bembrops greyi has the lowest average number of gill rakers of the nine Species and typically has a high number of vertebrae (30). Bembrops greyi has a short snout and a moderately long maxillary ten- tacle. The spinous dorsal fin pigmentation in Bembrops greyi is unique, in that, except for several small, irregular-shaped clear windows, the entire fin is jet-black (Fig. 1B). Bembrops greyi has three medium-length pyloric caeca, 6.8, 6.4, and 6.9 (left to right) % of SL. Distribution/Depth.—Eastern Atlantic Ocean in Gulf of Guinea off Grand-Bas- sam, Cote D’Ivoire, off Lagos, Nigeria; and from equator near Port Gentil, Gabon southward to Pointe Noire, Congo. We ex- amined three collections with Bembrops greyi taken with B. heterurus (Fig. 6). Ad- ditional records in Poll (1959) and Das & Nelson (1996). Our records of Bembrops greyi range from a depth of 101—494 m, with the largest series (19) containing many juveniles at the shallowest depth. Most col- lections are from 300—400 m. Poll (1959) reported it between 250—420 m, Blache et al. (1970) between 250—400 m, and Das & Nelson (1996) from 250—420 m. Material examined.—CU 48206 (19): 969 Geronimo 2-199; 01°26'S, 08°24’E; 400 m; 3 Sep 1963. CU 48215 (1): Geronimo 2- 220; 03°02’S, 09°21’E; 300 m; 6 Sep 1963. CU 48216 (6): Geronimo 2-205; 01°57’S, 08°47’E; 400 m; 4 Sep 1963. CU 48217 (1): Geronimo 2-238; 04°07'S, 10°23’E; 400 m; 8 Sep 1963. CU 48218 (1): Geronimo 2- 179; 00°02'S, 08°50’E; 295 m; 31 Aug 1963. CU 48219 (1): Geronimo 2-237; 04°03’S, 10°22’E; 300 m; 8 Sep 1963. CU 48220 (6): Geronimo 2-246; 04°31’S, 10°53’E; 300 m; 9 Sep 1963. CU 48221 (5): Geronimo 2-204; 01°56’S, 08°47’E; 300 m; 4 Sep 1963. UF 216947 (3): Pillsbury 51; 04°56'N, 05°01’W—04°56'N, 05°03'W; 329—494 m; 31 May 1964 [pyloric caeca dissection]. UF 220549 (19): Pillsbury 236; 05°20’N, 04°45’E—05°19'N, 04°48’E; 101— 128 m; 12 May 1965. Bembrops heterurus (Ribeiro, 1903) Robust duckbill Next to Bembrops macromma, B. heter- urus has the lowest number of dorsal soft rays. The dorsal fin formula for B. heteru- rus is VI, 14 to 16 soft rays, with a strong model count of 15. Anal fin rays are usually 18, only six specimens deviate from this count. Pectoral fin ray counts are either 26 or 27, and 27 is the more frequent number (Table 1). Bembrops heterurus has an intermediate number of lateral-line scales, between the high counts of B. anatirostris and B. go- bioides and the low counts of B. greyi and B. magnisquamis. Lateral-line scales range from 55 to 60, X = 56.5, being most similar to B. macromma (Table 2) and overlapping the counts of the three new species. Bembrops heterurus exceeds B. greyi in number of gill rakers; there are either four or five on upper limb, range from 13 to 16 on lower limb, and total counts range from 17 to 21 (Table 3). Bembrops heterurus and B. greyi differ markedly in number of ver- tebrae; B. heterurus typically has 28 where B. greyi typically has 30 vertebrae (Table 4). 970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON AFRICA ~ |bMossamedes Fig. 6. Three syntopic collections (*). Based on 50 specimens ranging from 80.7 to 214.5 mm SL, all have a snout greater than the eye; with none having the orbit length greater than or equal to length of snout (Tables 6 and 9). Maxillary tentacle lengths are quite sim- ilar in B. heterurus and B. greyi (Table 10), with B. heterurus having the third longest tentacle among the Atlantic species. The spinous dorsal fin pigmentation in Bembrops heterurus contrasts greatly with that described for B. greyi. Only the ante- Distribution of Bembrops greyi (A) and B. heterurus (@) based on specimens examined in this study. rior half of the first membrane is blackened in B. heterurus, with the rest of the fin clear or slightly dusky (Fig. 1D), whereas, except for small clear areas the entire fin is jet- black in B. greyi. Bembrops heterurus has three moderate- ly-long pyloric caeca, 9.7, 10.1, 9.6 (left to right) % of SL. Distribution/Depth.—Eastern Atlantic Ocean in Gulf of Guinea off southeastern Liberia; off Cote D’Ivoire in Bight of Benin and Bight of Bonny near Fernando Poo; VOLUME 111, NUMBER 4 southward along African coast from Port Gentil, Gabon to Pointe Noire, Congo; one record near Baia dos Tigres, southern An- gola (Fig. 6). We examined three collec- tions that had Bembrops heterurus taken with B. greyi. Bianchi et al. (1993) reported Bembrops heterurus as rare in Namibia its southernmost distribution along the west coast of Africa. Depth for Bembrops het- erurus used in this study ranged from 64— 494 m, with most records between 150 and 300 m. Poll (1959) reported this species be- tween 100 and 300 m. Fager & Longhurst (1968) found B. heterurus to be part of a “species group” centered around 200 m depth. Blache et al. (1990) gave a range be- tween 100—400 m. Das & Nelson (1996) reported a depth range for South American specimens of this species from 90—200 m. Material examined.—CU 48207 (6): Ge- ronimo 2-197; 01°30'S, 08°27’E; 200 m; 3 Sep 1963. CU 48208 (1): Geronimo 2-187; 00°32'S, 08°40’E; 300 m; 1 Sep 1963. CU 48209 (5): Geronimo 2-245; 04°31’S, 10°54’E; 200 m; 9 Sep 1963. CU 48210 (1): Geronimo 2-185; 00°32'S, 08°42’E; 200 m; 1 Sep 1963. CU 48211 (1): Geronimo 2- 213; 02°31’S, 08°51’E; 300 m; 5 Sep 1963. CU 48212 (5): Geronimo 2-212; 02°30’S, 08°58’E; 200 m; 5 Sep 1963. CU 48213 (2): Geronimo 2-227; 03°30'S, 08°53’E; 200 m; 7 Sep 1963. CU 48214 (5): Geronimo 2- 203; 02°01'S, 08°50'E; 200 m; 4 Sep 1963. TU 180287 (3): Undaunted 252; 16°41’S, 11°21’E; 164-182 m; 18 Mar 1968. UF 47051 (1): Geronimo 2-179; 00°02’S, 08°50'E; 295 m; 31 Aug 1963. UF 216954 (1): Pillsbury 51; 04°56'N, 05°01'’W— 04°56'30"N, 05°03'W; 329-494 m; 31 May 1964. UF 216958 (1): Pillsbury 82; 04°57'N, 09°30’ W—04°58'N, 09°32'W; 146-150 m; 5 Jun 1964. UF 216974 (4): Pillsbury 45; O5°05'N, 04°04'30"W-— 05°06'N, 04°06'W; 73-97 m; 30 May 1964. UF 220545 (7): Pillsbury 255; 03°49’N, 07°38'E—-03°48'N, 07°42'E; 264-269 m; 14 May 1965 [pyloric caeca dissection]. UF 220546 (8): Pillsbury 232; 05°56’N, 04°27'E-05°54'N, 04°27'E; 101-132 m; 11 971 May 1965. UF 220547 (5): Pillsbury 237; 05°19'N, 04°48’E-05°07'N, 04°55’E; 101 m; 12 May 1965. UF 220548 (2): Pillsbury 236; 05°20'N, 04°45’E-05°19'N, 04°48’E; 101-128 m; 12 May 1965. UF 220551 (7): Pillsbury 245; 04°32'N, 05°07'’E—04°31'N, 05°13’E; 64-119 m; 13 May 1965. UF 220552 (11): Pillsbury 254; 03°50'N, 07°08'’E—03°51'N, 07°12’E; 148-174 m; 14 May 1965. Bembrops macromma Ginsburg, 1955 Scaled-eye duckbill Dorsal fin rays in all specimens are VI, 14. Bembrops macromma is unique among the Atlantic species in its consistent low number of dorsal soft rays. The only other species with 14 soft dorsal rays are B. an- atirostris and B. heterurus and this is rare. Anal fin rays are usually 18 in number, but frequently are 17. Pectoral fin rays are usu- ally 25, but frequently 26, rarely 24 (Table 1). Bembrops macromma is similar to B. het- erurus in having an intermediate number of lateral-line scales, 53 to 60, X = 56.0 (Table 2). Gill rakers range from 4 to 6, usually 5, on upper limb, range from 12 to 15, usually 14 or 15, on lower limb, and total counts range from 16 to 21, predominantly 19 or 20 (Table 3). Bembrops macromma, like B. anatiros- tris and B. heterurus has a low number of vertebrae, usually 28, very rarely 27, the lowest vertebral number for Atlantic Bem- brops (Table 4). Both Bembrops macromma and B. het- erurus have stocky bodies, illustrated by the greater depth of head, depth of body at or- igin of first dorsal fin, and depth of caudal peduncle. Bembrops macromma has a short snout and large orbit whereas B. heterurus has a relatively long snout and a small orbit (Table 9). Bembrops macromma also differs from B. heterurus in length of maxillary tentacle, having the shortest tentacle of the nine species treated in this study (Table 10). Bembrops macromma is unique in the 972 genus in possessing scales on the dorsal surface of the eyes. Adults have 20—35 thin, cycloid scales covering each eye between the pupil and interorbit. Juveniles have few- er scales, but all specimens of this species possess these scales. In summary, Bembrops macromma has a low number of soft dorsal rays, usually 18 anal rays, and usually 25 pectoral rays. It has an intermediate number of lateral-line scales and a low number of vertebrae as does B. heterurus. Moreover, both B. ma- cromma and B. heterurus are relatively stout-bodied forms. However, B. macrom- ma has a shorter maxillary tentacle and also has a short snout and large orbit, whereas B. heterurus has a relatively long snout and a small orbit. Bembrops macromma has most of the anterior two membranes (rarely part of the third) blackened in the spinous dorsal fin (Fig. 1E) and B. heterurus has black pigmentation only in the anterior half of the first membrane. Bembrops macromma has three short py- loric caeca, 5.6, 6.0, and 6.0 (left to right) % of SL, across the dorsal surface of the stomach. Distribution/Depth.—Western Caribbean Sea on shelf along Honduras, Nicaragua, and Costa Rica; south side of Jamaica; north side of Cuba; three Bahama records one from just south of Grand Bahama Is- land, one from southwestern edge of Great Bahama Bank, and one at Great Inagua Is- land; west and northwest of Puerto Rico (taken with B. anatirostris); and from northern Leeward Islands (Fig. 5). We can- not confirm Das & Nelson’s (1996) Gulf of Mexico record (Fig. 12); this species ap- pears to be absent from the Gulf of Mexico. Uyeno et al. (1983) include B. macromma as one of two species in the family occur- ring off Suriname and French Guiana. We have not been able to substantiate this, and Das & Nelson (1996) also did not include South America in their range of this spe- cies. Specimens of Bembrops macromma used in this study ranged from 192-512 m in depth, with most specimens taken be- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tween 250 and 400 m. Ginsburg (1955) re- ported the types of this species from 274 and 549 m and Grey (1959) found it at 438 and 457 m. Das & Nelson (1996) reported it between 150 and 550 m, with “‘most specimens below 250 m’’. Material examined.—TU 12769 (2): Oregon 1341; 22°55'N, 79°16'W; 439 m; 16 Jul 1955. TU 12787 (1): Oregon 1341; 22°55'N, 79°16'W; 439 m; 16 Jul 1955. TU 18781 (3) and UF 202714 (5): Oregon 1879; 16°38'N, 81°39’W; 274 m; 22 Aug 1957 [pyloric caeca dissection]. TU 19933 (1): Oregon 1878; 16°39’N, 81°43’W; 230 m; 22 Aug 1957. TU 1801367 @)s@recom 1902; 11°27'N;, 83°11 We 247 iso see 1957. UF 15615 (2): Oresonmse2er 16°08’N, 81°13’W; 192-201 m; 6 Jun 1962. UF 207119 (2): Oregon 2649; 18°12’N, 64°18'W; 274 m; 6 Oct 1959. UF 220555 (1): Gerda 692; 26°35’N, 78°25'’W— 26°34'N, 78°26'W; 21 Jul 1965. USNM 157983 (2): Oregon “13445522 SOune 79°08'W; 366-411 m; 16 Jul 1955. USNM 304922 (4): Oregon 2658; 18°26’N, 67°11'30"W; 320 m; 7 Oct 1959. UF 47052 (2), USNM 304923 (8), USNM 304928 (2): Oregon 3549; 17°50'N, 77°52'W; 311 m; 16 May 1962. USNM 304925 (3), USNM 347204 (2): Oregon 10559; 23°04'N, 78°46'W; 347-377 m; 15 Dec 1969. USNM 304926 (5): Oregon 6699; 17°38'30’N, 62°16'W; 19 May 1967. USNM 304927 (3), USNM 342603 (2): Oregon 6700; 17°27'N, 62°04'W; 249-285 m; 19 May 1967. USNM 304929 (2): Oregon-II 10195; 14°17’N, 81°55'W; 329 m; 19 Nov 1968. USNM 342604 (1): Oregon 3625; 16°26’N, 81°35'W; 219 m; 6 Jun 1962. USNM 347205 (1): Oregon-II 10849; 20°50’N, 73°20'W; 311 m; 13 Dec 1969. CAS 61007 (2): Oregon-II 46095; 18°13'18’N, 67°19'30"W; 366 m; 21 Aug 1987. CAS 61010 (10): Oregon-II 46096; 18°13’'30"N, 67°18'42"W; 357-384 m; 21 Aug 1987. CAS 61011 (1): Oregon-II 46062; 18°30’00"N, 65°42'12"W; 441-512 m; 15 Aug 1987. VOLUME 111, NUMBER 4 Bembrops magnisquamis Ginsburg, 1955 Largescale duckbill The small number (8) of specimens pre- cludes much discussion and comparison. Dorsal fin formula is typically VI, 15 soft rays; anal fin has 17 rays; and pectoral fin ray counts are low, 22 to 24 (Table 1). Lat- eral-line scale counts are the lowest for any Atlantic Bembrops, ranging from 44 to 47 (Table 2). Gill raker counts are either 4 or 5 on upper limb, range from 13 to 16 on lower limb, and total counts range from 18 to 21 (Table 3). All eight specimens of Bembrops magnisquamis have 29 verte- brae, an intermediate number (Table 4). Bembrops magnisquamis has a relatively deep caudal peduncle and a very wide head (Table 8). Also, B. magnisquamis has a short snout and a large orbit, thus in three of the six specimens the orbit length is equal to length of snout and in the other three specimens the orbit length is greater than the snout (Table 9). Bembrops magnisquamis has a relatively long maxillary tentacle in comparison to the other eight species considered in this study. The anterior (first) membrane in the spi- nous dorsal fin is entirely black or with dark pigment at margin and base, with center of membrane clear. Membranes two and three are black on basal half and the pigmented basal portion is progressively less on suc- ceeding membranes four and five. Mem- brane six is clear in all specimens examined (Fig. 1F). This spinous dorsal fin pigmen- tation of B. magnisquamis is distinct but has slight resemblance to two of the new forms described below. There is a thin dash of pigment along the midline of the nape just anterior to the spinous dorsal fin in B. magnisquamis that is not present in any other Atlantic Bembrops. Distribution/Depth.—Type locality off S coast of Cuba; two sites in western Carib- bean Sea, one off Honduras, one off Costa Rica; one locality in eastern Caribbean Sea, S of Puerto Rico (Fig. 4); other records 973 (USNM) re-identified as B. macromma. We cannot confirm two records shown in Fig. 9, Das & Nelson (1996). Specimens of Bembrops magnisquamis used in this study ranged from 366—622 m. Ginsburg (1955) reported the types of the species from 366 and 465 m. Bullis & Struhsaker (1970) found B. magnisquamis between 368 and 640 m, with the highest densities in two depth strata, 368—457 m and 550-640 m. We examined one collection of B. magni- squamis taken with B. anatirostris off Costa Rica. Material examined.—USNM 45985 and 108395, holotype and paratype as listed in Ginsburg (1955). TU 180138 (1): Oregon 1883; 16°52’N, 81°30’W; 366 m; 23 Aug 1957. USNM 304924 (4): Oregon 3574; 12°31'N, 82°21’W; 366 m; 23 May 1962. CAS 61008 (1): Oregon-II 46026; 17°44’42"N, 66°12’42"W; 366-622 m; 9 Aug 1987. Bembrops ocellatus, new species Ocellate duckbill Rice? Bembrops anatirostris.—Das & Nelson, 1996 Gn part, locality in Caribbean Sea off Nicaragua). Material.—39 specimens (93.2—187.5 mm SL) from 23 localities. Holotype.—TU 181281, a female 135 mm SL, Atlantic Ocean, off Venezuela, 09°17'N, 59°19’W, 503 m, 4 Nov 1957, Oregon Sta. 1992, 45’ trawl. Paratypes.—TU_ 180353 (2): 117-126 mm SL, same data as holotype. TU 181668 (1): 91 mm SL, western Caribbean Sea, 16°41'N, 82°20’W, 549 m, 22 Aug 1957, Oregon Sta. 1872, 40’ flat trawl. CU 43877 (5): 97.0-146.0 mm SL, western Caribbean Sea, no locality data, May or Jun, 1962, Oregon Cruise 78, depth not known. UF 207128(1): 116 mm SL, eastern Caribbean Sea, off Venezuela, 11°36’N, 62°52’W, 394-421 m, 20 Apr 1960, Oregon Sta. 2780. USNM 307592 (3): 95.1-137.3 mm SL, Caribbean Sea off Nicaragua, 14°08’N, 974 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. 81°55'’W, 366-439 m, 21 May 1962, Oregon Sta. 3570. USNM 344486 (3): 112.6-128.8 mm SL, Caribbean Sea off Venezuela, 11°49’'N, 69°24'W, 549 m, 3 Oct 1963, Oregon Sta. 4412. USNM 347200 (1): 200 mm SL, Atlantic Ocean off Suri- name, 07°27’N, 54°30'W, 201 m, 16 May 1962, Oregon-II Sta. 10622. USNM 347201 (1): 125.2 mm SL, Atlantic Ocean off Gre- nada, 12°01'N, 61°53.5’W, 384—457 m, 26 Sep 1964, Oregon Sta. 5043. USNM 347202 (1): 142.1 mm SL, Caribbean Sea off Nicaragua, 12°26'N, 82°24’W, 503 m, 2 Jun 1962, Oregon Sta. 3609. USNM 347214 (1): 156.5 mm SL, Caribbean Sea off Nicaragua, 13°39'N, 81°52’W, 503 m, 13 Sep 1957, Oregon Sta. 1923. USNM 347215 (2): 145.2-159.7 mm SL, Carib- bean Sea off Colombia, 11°26’N, 74°14'W, 576 m, 3 Dec 1968, Oregon-II Sta. 10268. USNM 347216 (1): 169.4 mm SL, Atlantic Ocean N of St. Kitts & Nevis, 17°41'N, 62°50.5'W, 549-585 m, 18 May 1967, Oregon Sta. 6695. USNM 347217 (1): 144.0 mm SL, Atlantic Ocean NE off Dom- inica, 15°42'N, 61°08’W, 640 m, 1 Dec 1969, Oregon-II Sta. 10825. USNM 347254 (1): 143.0 mm SL, Caribbean Sea NW of Bembrops ocellatus, paratype, CAS 61006, 187.5 mm SL, female. Trinidad, 11°36'N, 62°42'W, 430 m, 19 Apr 1960, Oregon Sta. 2776. USNM 347261 (1): 93.2 mm SL, Caribbean Sea S of Ja- maica, 17°40'N, 77°55'W, 530 m, 16 May 1962, Oregon Sta. 3552. USNM 347262 (1): 93.5 mm SL, Caribbean Sea W of St. Kitts & Nevis, 17°20'N, 62°52'W, 549-567 m, 30 Sep 1964, Oregon Sta. 5072. CAS 14632 (1): 123.2 mm SL, Caribbean Sea off Venezuela, 11°53’N, 69°28’W, 421 m, 28 Sep 1963, Oregon Sta. 4408, 40’ flat trawl. CAS 61006 (1): 187.5 mm SL, Atlantic Ocean off NE Puerto Rico, 18°31'12’N, 65°40'36"W, 439-512 m, 15 Aug 1987, Oregon-II Sta. 46061. CAS 150896 (1): 118.9 mm SL, Atlantic Ocean off Surinam, 07°34'’N, 54°49'W, 411 m, 7 Nov 1957, Oregon 2007, 40’ flat trawl. CAS 98923 (1): 134.0 mm SL, Caribbean Sea, Lesser Antilles off St. Kitts and Nevis, 17°09'00"N, 66°44'48"W, 512-768 m, 4 Aug 1978, Oregon-II Sta. 24263. Other materials—USNM 347263 (1): 136.2 mm SL, Caribbean Sea W of An- guilla, 18°07'N, 63°20'W, 658 m, 7 Dec 1969, Oregon-II Sta. 10837. USNM 347264 (1): 140.4 mm SL, Caribbean Sea off Hon- duras, 16°51'N, 82°14’W, 585 m, 15 Nov VOLUME 111, NUMBER 4 1968, Oregon-II Sta. 10176. USNM 347272 (3): 141.7—210 mm SL, Caribbean Sea off Venezuela, 10°57’N, 67°02'’W, 457 m, 12 Oct 1963, Oregon Sta. 4454. USNM 347273 (3): 137.8-162.7 mm SL, SW Ca- ribbean Sea off NW coast of Colombia, 10°16.2'N, 75°54.5'W, 549 m, 25 May 1964, Oregon Sta. 4882. Diagnosis.—Dorsal fin usually VI-16 (rarely VI-15 or VI-17). Anal fin ranges from 16 to 19, usually 18. Pectoral fin usu- ally 26, but varies from 25 to 27. Lateral- line scales 55-65, most often 58 to 63, X = 60.2. Gill rakers usually 5 on upper limb, usually 15 on lower limb, and total count usually 20. Vertebral number usually 30 (29 in two). Ratio of head width divided by depth of head ranges from 1.3 to 1.6, usu- ally 1.4, X = 1.43. Snout short, usually slightly shorter, equal to or slightly longer than length of orbit (eye). Maxillary tenta- cle short, averaging only 2.6% of SL. The lateral-line has a moderately steep slope, reaching its lowest point under spines 5 or 6 of the first dorsal fin. Bembrops ocellatus possesses two (rarely four) wide black dor- sal saddles, located under the posterior por- tion of the second dorsal fin at rays 10-13, and on the posterior part of the caudal pe- duncle at the start of the secondary caudal fin rays, sharing this character only with B. quadrisella and B. raneyi in the Atlantic members of the genus, but differing from both B. quadrisella and B. raneyi in that the anterior two saddles, when present, are usu- ally less defined than the posterior two sad- dles. Description.—Bembrops ocellatus is most similar to B. gobioides, B. quadrisella, and B. raneyi, having high scale, fin-ray, and vertebral counts (Tables 1—4). Bem- brops ocellatus lacks the intense black, ‘x’ and ‘v’ secondary scale pigment that char- acterizes B. gobioides. Bembrops ocellatus lacks a black band in the anal fin, differing from B. anatirostris, B. heterurus, B. ma- cromma, and B. quadrisella whose males have a black band in the anal fin. Bembrops ocellatus, similar to B. raneyi, but differing 975 from all other Atlantic Bembrops, possesses a black “‘ocellus”’ spot at the dorsal base of the caudal fin in both sexes, although some males of this species have a basicaudal spot that is not an ocellus. Bembrops ocellatus has a black band in the distal part of the second dorsal fin and black at the base of rays 1—5 and 9—13, corresponding to where the dorsal body saddles are located. In larg- er specimens, the black forms two bands on the fin. This species lacks the elongate dor- sal spine found in male B. anatirostris and scaled eyes, unique to B. macromma. Bem- brops ocellatus never possesses vertically- oriented lateral markings, sometimes found in B. greyi, B. heterurus, and B. macromma. The first dorsal fin pigmentation is dis- tinct. There are two separate jet-black blotches, one at distal, anterior part of fin and a larger basal blotch. The wedge-shape, distal blotch is broadest across the first membrane, narrows progressively across second membrane, and its narrowed point extends halfway across the third membrane. The basal blotch broadens progressively from anterior, proximal third of first mem- brane to posterior half of third membrane, then the distal edge of blotch dips toward the base of the sixth spine, leaving most of the last membrane clear (Fig. 1G). Bembrops ocellatus tends to have a larg- er eye in relation to the snout with 82.2% of specimens having snout/eye ratio less than, or equal to, one; whereas B. quadri- sella has a somewhat longer snout with 93.1% of specimens having a snout/eye ra- tio greater than, or equal to, one. There is, however, considerable overlap. Distribution/Depth.—Atlantic Ocean off eastern coast of Venezuela (type locality); widely distributed in the Caribbean Sea: along the coast of Venezuela, off Costa Rica, Nicaragua, and Honduras, near Puerto Rico and along many islands of the Lesser Antilles (Fig. 8). We examined seven col- lections where Bembrops ocellatus was tak- en with B. quadrisella. Bembrops ocellatus is found from 394 to 549 m depth. Etymology.—The genus Bembrops is 976 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. (x). treated as masculine; the species name ocel- latus is from the Latin word “‘ocellus”’, meaning a little eye (Jaeger 1966) referring to the black spot on the dorsal base of the caudal fin. Bembrops quadrisella, new species Saddleback duckbill Fig. 9 Bembrops anatirostris.—Das & Nelson, 1996 (in part, locality in Caribbean Sea off Nicaragua). Bembrops gobioides.—Das & Nelson, 1996 (in part, locality in Lesser Antilles). Material.—123 specimens (101.2—231.0 mm SL) from 32 localities. Holotype-—TU 180351, a female 195 mm in SL, Atlantic Ocean, off Suriname, 07°38’N, 54°43’W, 457 m, 7 Nov 1957, Oregon Sta. 2008, 40’ trawl, bottom listed as “‘blue mud’’. Paratypes.—TU 181667 (1): 178 mm SL, same data as holotype. TU 180350 (1): 159 mm SL, Atlantic Ocean, off Suriname, 07°34'N, 54°49’W, 411 m, 7 Nov 1957, Oregon Sta. 2007. TU 180259 (1): 127 mm SL, western Caribbean Sea, off Nicaragua, 16°39'N, 82°26'’W, 457 m, 22 Aug 1957, Oregon Sta. 1871. TU 180353 (1): 145 mm SL, western Caribbean Sea, 16°41'N, Distribution of Bembrops ocellatus (@) based on specimens examined in this study. Type locality 82°20'W, 549 m, 22 Aug 1957, Oregon Sta. 1872. TU 20037 (1): 185 mm’ SEveastem Caribbean Sea, 11°35’N, 62°41'W, 388—457 m, 23 Sep 1958; Oregon StasiZ3555 CU) 43863 (1): 227 mm SL, western Caribbean Sea, no locality data, May or Jun 1962, Oregon Cruise 78. CU 77466 (1): 163 mm SL, western Caribbean Sea, no locality data, May or Jun 1962, Oregon Cruise 78. USNM 304913 (4): 127.2—211 mm SL, Ca- ribbean Sea off Nicaragua, 12°26’N, 82°24'W, 503 m, 2 Jun 1962, Oregon Sta. 3609. USNM 304917 (2): 192—235 mm SL, Atlantic Ocean NE of Dominica, 15°39'N, 61°10'W, 649 m, 5 Mar 1966, Oregon Sta. 5929. USNM 347206 (4): 139.8—168.0 mm SL, Caribbean Sea off Nicaragua, 14°10'N, 81°55'W, 439-457 m, 21 May 1962, Oregon Sta. 3565. USNM 347207 (3): 154.2-162.6 mm SL, Caribbean Sea off Honduras, 14°10’N, 81°50’W, 549-604 m, 21 May 1962, Oregon Sta. 3571. USNM 347208 and 347258 (2): 182—231 mm SL, Lesser Antilles, 17°06'N, 62°17'W, 589 m, 8 Dec 1969, Oregon-II Sta. 10843. USNM 347209 (1): 222 mm SL, Atlantic Ocean off Dominica, 15°36’N, 61°13’W, 503 m, 4 Mar 1966, Oregon Sta. 5926. USNM 347210 (2): 173-205 mm SL, Caribbean Sea E of VOLUME 111, NUMBER 4 977 Fig. 9. Bembrops quadrisella, paratype, CAS 61004, 197.5 mm SL, male. St. Croix, 17°42'N, 63°58'W, 741 m, 3 Dec 1969, Oregon-II Sta. 10832. USNM 347211 (2): 200-201 mm SL, eastern Caribbean Sea, 11°40’N, 62°33’W, 585-621 m, 24 Sep 1964, Oregon Sta. 5039. USNM 347218 (11): 101.2-189.5 mm SL, Caribbean Sea off Costa Rica, 12°25’N, 82°15'W, 549-585 m, 23 May 1962, Oregon Sta. 3576. USNM 347219 (1): 192.5 mm SL, Caribbean Sea off Venezuela, 11°53’N, 69°28'W, 421 m, 28 Sep 1963, Oregon Sta. 4408. USNM 347220 (6): 170.4—228 mm SL, Atlantic Ocean NE off Dominica, 15°42’N, 61°08'’W, 640 m, 1 Dec 1969, Oregon-II Sta. 10825. USNM 347221 and 347256 (3): 176.0—230 mm SL, Atlantic Ocean N of St. Kitts & Nevis, 17°41'N, 62°50.5’W, 549— 585 m, 18 May 1967, Oregon Sta. 6695. USNM 347255 (1): 215 mm SL, Caribbean Sea E of St. Croix, 17°36'N, 63°32’W, 439— 476 m, 30 Sep 1959, Oregon Sta. 2636. USNM 347257 (1): 204 mm SL, Atlantic Ocean NE of St. Croix, 18°18’'N, 63°24’W, 658 m, 10 Dec 1969, Oregon-II Sta. 10847. USNM 347259 (1): 149.2 mm SL, Carib- bean Sea off Venezuela, 10°54’N, 67°08’W, 402 m, 12 Oct 1963, Oregon 4453. USNM 347260 (11): 113.1-190.5 mm SL, Carib- bean Sea off Honduras, 16°58’N, 87°53’W, 457-732 m, 10 Jun 1962, Oregon Sta. 3635. USNM 347266 (11): 130.0—-188.0 mm SL, Caribbean Sea off Honduras, 16°51'N, 82°14'’W, 585 m, 15 Nov 1968, Oregon-II Sta. 10176. USNM 347267 (10): 132.2—-162.0 mm SL, Caribbean Sea off Costa Rica, 12°35'N, 82°19'W, 457 m, 23 May 1962, Oregon Sta. 3575. USNM 347268 (5): 130.9-158.1 mm SL, Carib- bean Sea off Honduras, 16°44’N, 87°55’W, 347 m, 9 Jun 1962, Oregon Sta. 3634. USNM 347269 (8): 120.0—223 mm SL, Ca- ribbean Sea SE of Puerto Rico, 17°40'N, 63°40'W, 658-695 m, 30 Sep 1959, Oregon Sta. 2637. CAS 56879 (1): 205 mm SL, Ca- ribbean Sea, Lesser Antilles off St. Kitts and Nevis, 17°09'00"N, 66°44’48’W, 512-— 768 m, 4 Aug 1978, Oregon-II Sta. 24263. CAS 61004 (2): 134.0-197.5 mm SL, At- 978 lantic Ocean, U.S. Virgin Islands, N of St. Thomas Island, 18°32’42"N, 65°42’00"W, 732-777 m, 16 Aug 1987, Oregon-II Sta. 46067, 65’ trawl. CAS 61005 (1): 205 mm SL, Atlantic Ocean, Puerto Rico, NW coast off Bahir de Aquadilla, 18°26'42’N, 67°14'42"W, 658-668 m, 21 Aug 1987, Oregon-II Sta. 46094. Other materials —USNM 347270 (3): 147.1—192 mm SL, Caribbean Sea off Ven- ezuela, 11°10'N, 68°08’W, 402 m, 11 Oct 1963, Oregon Sta. 4451. USNM 347271 (17): 114.7-184 mm SL, Caribbean Sea off Nicaragua, 17°02'N, 81°27'W, 549-585 m, 7 Jun 1962, Oregon Sta. 3629. CAS 47193 (1): 175 mm SL, Caribbean Sea off Vene- zuela, 12°48’N, 70°10'W, 366-914 m, 26 Feb 1973, Oregon-II Sta. 129. CAS 56927 (1): 188 mm SL, Caribbean Sea, Lesser An- tilles off St. Kitts and Nevis, 17°15'54’N, 62°16'48"W, 567-640 m, 5 Aug 1978, Oregon-II Sta. 24265. Diagnosis.—Dorsal fin usually VI-16 (82% of specimens). Anal fin 17 or 18, rarely 16 or 19. Pectoral rays usually 26 or 27, sometimes 25 or 28. Lateral-line scales 57-65, X = 61.1. Gill rakers always 5 on upper limb, usually 15 or 16 on lower limb, and usually a total count of 20 or 21. Ver- tebral number usually 30 (29 in six). Ratio of head width divided by depth of head ranges from 1.2 to 1.5, usually 1.4, X = 1.39. Upper and lower jaw moderately long, and postorbital distance also moderately long. Snout short, equal to or slightly great- er than length of orbit (eye). Maxillary ten- tacle short, averaging only 2.1% SL. The lateral-line has a gentle slope, descending gradually to the straight portion under the space between the two dorsal fins or under the anterior rays of the second dorsal fin. Benbrops quadrisella possesses four wide black dorsal saddles, located at the origin of the spinous dorsal fin, under rays 2—6 of the second dorsal fin, under rays 10—13 of the second dorsal fin, and on the posterior part of the caudal peduncle at the start of the secondary caudal fin rays, sharing this character with B. raneyi. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Description.—Bembrops quadrisella is most similar to B. gobioides, B. ocellatus, and B. raneyi, having high scale, fin-ray, and vertebral counts (Tables 1—4). Bem- brops quadrisella lacks the intensive black ‘x’ and ‘v’ secondary scale pigment that characterizes B. gobioides. Male B. quad- risella have the distal % to % of the anal fin black, similar to B. anatirostris, B. heteru- rus, and B. macromma; other Atlantic Bem- brops, B. greyi, B. gobioides, B. magni- squamis, B. ocellatus, and B. raneyi have clear or slightly dusky fins, without broad black bands in either sex. Bembrops quad- risella is the only Atlantic species in the genus possessing white pigment in the spi- nous and second dorsal, anal, and pectoral fins. Female B. qguadrisella possess a black *“‘ocellus”’ spot at the dorsal base of the cau- dal fin, differing from B. anatirostris, B. heterurus, B. macromma, and B. magni- squamis that lack ocelli in both sexes, but is similar to B. greyi and B. gobioides, where females also possess an ocellus, but differing from B. ocellatus and B. raneyi that have a basicaudal spot in both sexes. Bembrops quadrisella lacks black bands in the second dorsal fin, having small intense black marks only at the base of several an- terior rays. This species lacks the elongate spine found on male B. anatirostris and scaled eyes, unique to B. macromma. Bem- brops quadrisella never possesses vertical- ly-oriented lateral blotches, sometimes found in B. greyi, B. heterurus, and B. ma- cromma. The first dorsal fin pigmentation of fe- male B. quadrisella is distinct (Fig. 1H). There is a small spot of black at base of first membrane and a black, submarginal band, broader on first two membranes and then a narrowed band extending across third, fourth, fifth, and sixth membranes. The greater part of the fin is either clear or lightly sprinkled with melanaphores. Male B. quadrisella usually have a first dorsal fin pattern similar to the female, but some specimens lack this distinctive pattern, pos- VOLUME 111, NUMBER 4 mE 7) Ss ISPANIOLA 979 pee ieee Fee a o Fig. 10. Distribution of Bembrops quadrisella (@) based on specimens examined in this study. Type locality (%). sessing only a light wash of melanophores over the entire fin. Distribution/Depth.—Atlantic Ocean off Suriname (type locality; second site near type locality; widely distributed in the Ca- ribbean Sea: along the coast of Venezuela; off Costa Rica, Nicaragua, and Honduras; S of Jamaica; near Puerto Rico and along length of Lesser Antilles (Fig. 10). We ex- amined seven collections where Bembrops quadrisella was taken with B. ocellatus and one collection taken with B. anatirostris. Bembrops quadrisella is found from 347 to 914 m, usually between 400 and 600 m. Etymology.—The species name quadri- sella is from the Latin “‘quadrus”’, meaning fourfold or four, and “‘sella’’, a seat or sad- dle (Jaeger 1966) in reference to the four well-developed saddles on the dorsum of the body. Bembrops raneyi, new species Bahama duckbill Fig. 11 Material.—Twenty nine specimens (69.0-176.0 mm SL) from ten localities. Holotype.—TU 180135, a male 136 mm SL, Atlantic Ocean, just east of Cay Sal Bank, 23°59’N, 79°43’'W, 640 m, 24 Jul 1957; Combat Sta. 450; 10’ beam trawl. Paratypes.—CU 77467 (1): 110 mm SL, TU 16849 (6): 69.0-145.0 mm SL; same data as holotype. TU 180260 (4): 107.0— 164.0 mm SL; Atlantic Ocean, E of Vero Beach, Florida and north of Matanilla Shoal light; 27°53’N, 79°09’W; 686-759 m; 9 Jun 1958; Silver Bay Sta. 442; 40’ flat trawl. UF 212111 (1): 137.4 mm SL; Atlantic Ocean, Bahama Islands, Elbow Bank, W of Cat Cay, 26°27'N, 79°21'’W-26°36'N, 79°24'W; 531-540 m; 25 Jun 1963; Gerda Sta. 158; otter trawl. UF 217644 (1): 125.1 mm SL; Atlantic Ocean, Bahama Islands, Elbow Bank, W of Alice Town, Bimini Is- land, 25°49’N, 79°21’W—25°53’N, 79°22'W; 284-293 m; | Jul 1965; Gerda Sta. 646. UF 220553 (1): 167.3 mm SL; Atlantic Ocean, Grand Bahama Island, S of Freeport, 26°26'N, 78°39’ W-—26°27'N, 78°41'W; 724 m; 22 Jul 1965; Gerda Sta. 709. UF 220554 (1): 109.9 mm SL, Atlantic Ocean, Bahama Islands, NW of Berry Islands, 25°56'N, 78°09’ to 05’W; 595-711 m; 20 Jul 1965; Gerda Sta. 679. USNM 269552 (4): 93.0— 146.8 mm SL and one cleared and stained, 115.0 mm SL; Atlantic Ocean, off Nassau, exact locality uncertain, probably 26 Oct 1961; Silver Bay Sta. 3479. USNM 347223 (5): 82.9-151.7 mm SL; Atlantic Ocean in Straits of Florida, 23°09'N, 80°08’W; 466 980 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 11. m; 16 Dec 1969; Oregon-II Sta. 10863. USNM 347252 (3): 104.4-176.0 mm SL; Atlantic Ocean in Straits of Florida; 23°59'N, 79°17'W; 530-549 m; 17 Nov 1960; Silver Bay Sta. 2469. USNM 347253 (1): 117.0 mm SL; Atlantic Ocean in Straits of Florida, 23°40’N, 79°18’W; 530 m; 5 Nov 1960; Silver Bay Sta. 2458. Diagnosis.—Dorsal fin usually VI-16 (VI-15 in three, VI-17 in one). Anal fin usu- ally 18 (17 in nine). Pectoral fin 25-29, usually 27. Lateral-line scales 56—61, xe— 59.0. Gill rakers usually 5 on upper limb, 14 on lower limb, and total number usually 19. Vertebral number usually 30 (29 in two). Maxillary tentacle a long triangle, av- eraging 4.1% SL. Head very broad, width averages 1.5 times its depth. Snout short, slightly longer or shorter than orbit. Upper, lower jaws and postorbital lengths moder- ately short. The lateral-line has a relatively gentle slope reaching its lowest point be- tween the two dorsal fins. Body pale. First dorsal fin with two black blotches, the smaller, a narrow submarginal bar spanning the first two membranes, the larger extend- ing along base of fin, crossing the first to fourth membranes. The interspace between Bembrops raneyi, paratype, USNM 347223; 141.7 mm SL, male. the distal and basal blotches is clear (Fig. 11). Bembrops raneyi possesses four wide dark dorsal saddles, located at the origin of the spinous dorsal fin, under rays 2—6 of the second dorsal fin, under rays 10—13 of sec- ond dorsal fin, and on the posterior part of the caudal peduncle at the start of the sec- ondary caudal fin rays, sharing this char- acter with only B. quadrisella. Description.—Bembrops raneyi is most similar to B. gobioides, B. ocellatus, and B. quadrisella, having high scale, fin-ray, and vertebral counts (Tables 1—4). Bembrops raneyi lacks the dark black ‘x’ and ‘v’ sec- ondary scale pigment that characterizes B. gobioides and B. greyi. Bembrops raneyi lacks both scaled eyes as found in B. ma- cromma and elongate second dorsal spine of B. anatirostris. B. raneyi lacks black bands in the anal fin, distinguishing it from B. anatirostris, B. heterurus, B. macromma, and B. quadrisella whose males possess black in the anal fin. Both sexes of B. ra- neyi have a dorsally-placed, basicaudal spot; the only other Atlantic Bembrops sharing this character is B. ocellatus. Bem- brops raneyi never possesses vertically-ori- VOLUME 111, NUMBER 4 a, : 5 Fig. 12. ented lateral markings as sometimes found in B. greyi, B. heterurus, and B. macromma. Bembrops raneyi has three very short py- loric caeca, 2.6, 3.9, and 3.4 (left to right) % of SL, across the stomach. These are the shortest caeca of any Atlantic species ex- amined in the genus Bembrops. Bembrops raneyi has the longest tentacle for any At- lantic species of Bembrops except for B. anatirostris (Table 10). Distribution/Depth.—Atlantic Ocean in western, southwestern and northwestern Bahamas Islands; (Fig. 12), three species of Distribution of Bembrops raneyi (@) based on specimens examined in this study. Type locality (%*). Bembrops: B. anatirostris, B. macromma, and B. gobioides have been taken in area near records of B. raneyi. Bembrops raneyi is one of the deeper dwelling species in the genus, being found from 284 to 759 m, with most specimens taken below 500 m. Etymology.—We take great pleasure in naming this species after the late Dr. Ed- ward C. Raney, former Professor of Ichthy- ology at Cornell University and teacher to many of North America’s ichthyologists, our mentor, friend, and professional col- league, who had a long-standing interest in 982 percophids and organized much of the study materials we have used in the present research. We made use of his notes, and an early draft of a manuscript describing the species that now bears his name. Discussion Our counts and measurements agreed, for the most part, with those of Ginsburg (1955). Our interpretation of the structural bases of the fin rays was verified by nu- merous x-rays. Our counts of 113 B. ana- tirostris resulted in eight with 14 dorsal rays and 105 with 15 dorsal rays. Bembrops anatirostris and B. gobioides have the highest lateral-line scale counts of the nine species of Bembrops from the At- lantic Ocean. Both males and females dis- play a silvery sheen on the prepectoral area and for a varying distance forward on side of isthmus. None of the other seven species was observed to have such coloration. Bem- brops anatirostris and B. gobioides differ, however, in several characteristics. Previous studies have noted the elongate dorsal spine of B. anatirostris. Ginsburg (1955) and Grey (1959) stated that the second dorsal spine of male was prolonged, however Das & Nelson (1996) stated that the second or third spine of the male was produced. We found no male B. anatirostris with pro- longed third dorsal spine, all were second dorsal spines. Bembrops anatirostris usu- ally has 15 dorsal rays whereas B. gobioi- des usually has 17 dorsal rays; B. anatiros- tris invariably has 28 vertebrae whereas B. gobioides usually has 30. Bembrops anati- rostris has a greater head, and caudal pe- duncle depth than B. gobioides. Bembrops anatirostris has a greater head length, head width, snout length, jaw length, and post- orbital length than B. gobioides. Bembrops anatirostris has a smaller orbit relative to snout length and a very long attenuate max- illary tentacle in comparison to B. gobioi- des which has a larger orbit that is some- times equal to snout length, especially in small juveniles and has a maxillary tentacle PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of moderate length. In addition to these dif- ferences, there are a number of differences in pigmentation of fins and body that are described above in the species accounts. Bembrops macromma and B. magni- squamis were the only two previously known western north Atlantic forms pos- sessing medium to large scales. These two forms differ markedly in a number of ways. Unfortunately, we have only eight speci- mens of B. magnisquamis to compare with B. macromma. Bembrops macromma in- variably has 14 dorsal rays, usually 18 anal rays, and usually 25 pectoral rays, whereas B. magnisquamis has usually 15, occasion- ally 16 dorsal rays, 17 anal rays and either 23 or 24 pectoral rays. The number of lat- eral-line scales range from 53 to 61 in B. macromma Vs. 44 to 47 in B. magnisquam- is. Bembrops macromma usually has 28 vertebrae, occasionally 27 whereas B. mag- nisquamis has 29. Bembrops macromma has a greater head and body depth, but a similar caudal peduncle depth as B. mag- nisquamis. Bembrops macromma has the shortest maxillary tentacle of the nine spe- cies of Bembrops included in this study in contrast to B. magnisquamis which has a moderately long maxillary tentacle. Pig- mentation differences are described in de- tail in the species accounts. Bembrops ocellatus, B. quadrisella, and B. raneyi are quite similar in most meristics and morphometrics. These three new spe- cies typically have 16 dorsal rays and 18 anal rays, but occasionally have 15 of the former and 17 of the latter. Bembrops ocel- latus and B. quadrisella typically have 26 pectoral rays whereas B. raneyi usually has 27 pectoral rays. Bembrops ocellatus and B. quadrisella have more lateral-line scales, 60 to 63 in former and 59 to 64 in the latter; B. raneyi with a lower count, 56 to 61. All three species usually have five gill rakers on upper limb, but B. ocellatus and B. quadrisella usually have 15 rakers on lower limb and B. raneyi usually has 14 rakers on lower limb, so total gill raker counts usually are 20 for B. ocellatus and B. quadrisella VOLUME 111, NUMBER 4 but 19 for B. raneyi. Bembrops raneyi has slightly greater body depth, less head depth, greater head width, and greater orbit length relative to snout length than the other two species. Bembrops quadrisella has a greater head length, greater head depth, greater snout length, greater upper jaw, lower jaw, and postorbital lengths than either B. ocel- latus or B. raneyi. Bembrops raneyi has the longest maxillary tentacle, B. ocellatus the next longest, and B. guadrisella the shortest maxillary tentacle of the three new species. The maxillary tentacle of B. raneyi is slightly shorter than that of B. anatirostris, and the tentacle of B. quadrisella is only slightly longer than that of B. macromma. There are some distinct pigmentation pat- terns of fins and body but these are de- scribed above in detail. We discuss B. greyi and B. heterurus to- gether because our samples of both species are from the eastern Atlantic and both spe- cies have a general appearance distinct from the seven western north Atlantic species. These two species are quite easily separat- ed: Bembrops greyi usually has 16 dorsal rays, B. heterurus usually has 15; B. greyi usually has either 17 or 18 anal rays, B. heterurus typically has 18. Bembrops greyi usually has 26 pectoral rays, whereas: B. heterurus usually has 27. Bembrops greyi has a range in lateral-line scale counts from 45 to 52, whereas B. heterurus ranges from 55 to 60. There are fewer gill rakers in B. greyi than in B. heterurus. Bembrops greyi typicaily has 30 vertebrae whereas B. het- erurus usually has 28. Das & Nelson (1996) reported B. heterurus from the western At- lantic with 17 anal rays (84% of speci- mens), differing from our findings from eastern Atlantic specimens with specimens having 18 anal rays (92% of specimens). Das & Nelson’s (1996) analysis of western south Atlantic Bembrops heterurus shows that nearly all proportions of this species differed from our observations on the east- ern Atlantic populations (considered B. het- erurus by Poll (1959)), suggesting that the 983 eastern Atlantic form is an undescribed spe- cies. Bembrops heterurus has a greater body depth, greater head depth, greater head width, greater interorbital width, and great- er snout length than B. greyi. Bembrops greyi has a greater head length and greater orbit length than B. heterurus. The relative length of snout versus length of orbit re- veals that the orbital horizontal distance is usually greater than the snout length in B. greyi but never greater than or equal to snout length in B. heterurus. The maxillary tentacle is of moderate size in both B. greyi and heterurus. There has been little discussion concern- ing sexual dimorphism in the Atlantic members of the genus Bembrops. Ginsburg (1955) noted the prolongation of the second dorsal spine in “larger males” of B. ana- tirostris. Grey (1959) also noted this sexual dimorphism in B. anatirostris and suggest- ed “it is possible that other species may also exhibit sexual dimorphism’’, but pro- vided no further observations. Ginsburg (1955:631) noted “the smaller specimens (referring to the genus Bembrops) having a spot on the caudal near its base, nearer to upper than lower margin, disappearing with growth ...’. Our observations on series of the Atlantic species of Bembrops shows this statement to be partially correct. All small specimens of B. gobioides and B. greyi pos- sess an ocellus as described by Ginsburg (1955), however this mark is retained in adult females, but faded or completely lost in adult males. Bembrops quadrisella is also sexually dimorphic in females having a caudal spot that the males lack. Masuda et al. (1984) reported similar sexual dimor- phism with the Pacific species, B. caudi- macula and B. filifera. Two Atlantic spe- cies, B. ocellatus and B. raneyi and one Pa- cific species, B. curvatura have both sexes possessing a caudal spot. We find Bembrops anatirostris, B. heterurus, B. macromma, and B. magnisquamis lack a caudal spot for all life history stages. Das & Nelson (1996) reported that certain specimens of Bem- 984 brops species possess a caudal spot, but did not relate it to the sex of the specimen. Bembrops anatirostris, B. macromma, and B. quadrisella are sexually dimorphic in anal fin pigment, with males having black in the fin that is lacking in females. All species of Bembrops examined in this study are sexually dimorphic in genital pa- pilla size; males having a longer and more robust, papilla, females having a much small- er papilla. This dimorphism in papilla sizes reaches an extreme in the Pacific species, Bembrops curvatura, whose males possess an elongate, club-shaped papilla, 2—3 times the length of the papilla of the female. Most Atlantic Bembrops have a gentle slope to the lateral-line, descending to its lowest point on the body between the two dorsal fins. Interspecific variation of this character ranges from B. heterurus and B. macromma where the slope is so gradual that the straight section of the lateral-line starts under the anterior part of the second dorsal fin to B. gobioides that possesses an abrupt slope where the lateral-line descends to the straight section under the first dorsal fin. Among the new species, Bembrops ocel- latus and B. raneyi have slightly steeper slopes to the lateral-line than B. quadrisella. The nine species of Atlantic Bembrops have 27 to 30 vertebrae. Miller & Jurgen- son (1973) listed precaudal, caudal and total counts for these species, but their values for B. macromma are partially in error. They report Bembrops normally having nine pre- caudal vertebrae, agreeing with our findings (except for an infrequent count of ten in B. anatirostris). Vertebral number seems to be a conservative character with little variation within most species (Table 4). What varia- tion in vertebral number, within and be- tween species, that does exist, results from differences in caudal vertebrae. This differs from Chrionema, the sister group to Bem- brops (sensu Nelson 1994) since Iwamoto & Steiger (1976) report 19 caudal vertebrae in Chrionema and either 8 or 9 precaudal vertebrae for a total of 27 or 28. We cannot agree with Das & Nelson’s PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (1996) statement in the description of the genus that species of Bembrops possess “two small spinous processes on the pos- terior tip of posttemporal’’. In five species examined (Fig. 3) there is only a single prominent spine projecting dorso-posteri- orly from the rear of the posttemporal bone. Just posterior and lateral to the posttempor- al spine, a protuberance on the supracleith- rum frequently projects through the epider- mis suggesting the presence of a second spine near the anterior keeled lateral-line scale. Shape, size, and degree of curvature appears to vary among the species. Acknowledgments We are most grateful for Dr. Raney’s per- sonal notes, recorded data, personal corre- spondence, x-ray files, etc., with regards to the systematics and life history of perco- phids. Among his files was an English translation of a critical paper (Poll 1959) in French, for which we are most appreciative. Raney received assistance from Frederick H. Berry, Harvey Bullis, Bruce B. Collette, Robert E. Jenkins, Leslie W. Knapp, the late Ermest A. Lachner, Giles Mead, Bill Richards, C. Richard Robins, Stewart Springer, Frank T. Williams, and perhaps others, all to whom we extend our hearty thanks. We extend our sincere appreciation to Jo- seph S. Nelson for discussions with regards to percophids. We thank Nancy Voss for valuable exploratory station data and for in- formation with regards to specimens formerly housed at University of Miami (UMML). We thank C. R. Robins for the transfer of mate- rials formerly on loan to Cornell University (CU) to Tulane University (TU). We thank Amy McCune for specimens from Cornell University, Susan Jewett and David Smith for the loan of specimens and assistance with the holdings of percophids at the National Museum of Natural History (USNM), and John McCosker and David Catania for specimens from the California Academy of Sciences (CAS). We thank George Burgess for printouts of holdings at VOLUME 111, NUMBER 4 the University of Florida (UF) and for other information with regards to the transfer of UMML collections to the University of Florida. We thank Henry L. Bart and Carter R. Gilbert for permission to make dissections on specimens under their care. We are grateful to former Tulane Uni- versity students, John H. Caruso, Michael Dahlberg, and Edwin Joseph for their spe- cial efforts in obtaining Bembrops speci- mens while on board the M/V Oregon. We thank Kenneth Kitchell and Stephen Schierling of the Louisiana State University Foreign Languages Department for advice on the Latin derivation of the names of two of the new species. Also, we thank Henry L. Bart and his staff, especially Michael Taylor, for the many courtesies with regards to working space, handling of loans, cataloging speci- mens, and retrieval of cataloged data at Tu- lane University. Contribution number LSU-CFI-97-06 of the LSU Center for Coastal, Energy, and Environmental Resources, Coastal Fisheries Institute, Baton Roue, LA 70803-7503. Literature Cited Bianchi, G., K. E. Carpenter, J.-P. Roux, FE J. Molloy, D. Boyer, & H. J. Boyer. 1993. FAO species identification field guide for fishery purposes. The living resources of Namibia. Food and Ag- riculture Organization of the United Nations, Rome, 250 pp. Blache, J., J. Cadenat, & A. Stauch. 1970. Clés de détermination des poissons de mer signalés dans L Atlantique oriental. Faune Tropicale. XVIII. O.R.S.T.O.M, Paris, 479 pp. Boschung, H. T. 1992. Catalogue of freshwater and marine fishes of Alabama.—Bulletin Alabama Museum of Natural History 14:1—266. Bullis, H. R., & P J. Struhsaker. 1970. Fish fauna of the western Caribbean upper slope.—Quarterly Journal of the Florida Academy of Sciences 33(1):43-76. Das, M. K., & J. S. Nelson. 1996. Revision of the Percophid genus Bembrops (Actinopterygii: Perciformes).—Bulletin of Marine Science 59(1):9—44. Fager, E. W., & A. R. Longhurst. 1968. Recurrent group analysis of species assemblages of de- 985 mersal fish in the Gulf of Guinea.—Journal of the Fisheries Research Board of Canada 25(7): 1405-1421. Ginsburg, I. 1955. Fishes of the family Percophididae from the coasts of eastern United States and the West Indies with descriptions of four new spe- cies.—Proceedings of the United States Nation- al Museum 104(3347):623—639. Grey, M. 1959. Deep sea fishes from the Gulf of Mex- ico with the description of a new species.—Fiel- diana-Zoology 39(29):323-346. Hubbs, C. L., & K. F Lagler. 1958. Fishes of the Great Lakes region. Cranbrook Institute of Science, Bulletin 26, 213 pp. , & . 1964. Fishes of the Great Lakes region. University of Michigan Press, Ann Ar- bor, Michigan, 213 pp. Iwamoto, T., & J. C. Steiger. 1976. Percophidid fishes of the genus Chrionema Gilbert.—Bulletin of Marine Sciences 26(4):488—498. Jaeger, E. C. 1966. A source-book of biological names and terms. C. C. Thomas, Springfield, Illinois, 323 pp. Masuda, H., K. Amaoka, C. Araga, T. Uyeno, & T. Yoshina. 1984. The fishes of the Japanese Ar- chipelago. Tokai University Press, Tokyo, 437 Pp. Miller, G. L., & S. C. Jorgenson. 1973. Meristic char- acters of some marine fishes of the western At- lantic Ocean.—Fishery Bulletin 71(1):301-312. Nelson, J. S. 1976. Fishes of the World. Wiley-Inter- science, New York, 416 pp. . 1978. Bembrops morelandi, a new percophi- did fish from New Zealand, with notes on other members of the genus.—National Museum of New Zealand Records 1(14):237—241. Nelson, J. S. 1984. Fishes of the World, 2nd Ed. Wi- ley-Interscience Publication, New York, 523 pp. Nelson, J. S. 1994. Fishes of the World, 3rd Ed. John Wiley & Sons, New York, 600 pp. Poll, M. 1959. Poissons IV. Teleosteens Acanthopter- giens (Deoxieme Partie). Expédition Océano- graphique Belge dans les Eaux Cotieres Africaines de L Atlantique Sud (1948-1949) 4(3B):1—417. Richards, W. J. 1990. List of the fishes of the western central Atlantic and the status of early life stage information. NOAA Technical Memorandum NMEFS-SEFC-267, 88 pp. Suzuki, T., & T. Nakabo. 1996. Revision of the genus Acanthaphritis (Percophidae) with the descrip- tion of a new species.—Ichthyological Research 43:441—454. Uyeno, T., K. Matsuura, & E. Fujii. 1983. Fishes trawled off Suriname and French Guiana. Japan Marine Fishery Resource Research Center, To- kyo, 519 pp. Williams, F T. 1968. Report on the Guinean Trawling Survey. 3 vols. Organization of African Unity- Scientific, Technical and Research Commission, Lagos, Nigeria. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):986—-991. 1998. A new species of cardinalfish (Apogonidae) from the Philippines, with comments on species of Apogon with six first dorsal spines Thomas H. Fraser W. Dexter Bender and Associates, Inc., 2052 Virginia Avenue, Fort Myers, Florida 33901, U.S.A. Abstract.—A new species of Apogon (A. bryx) from deep water is described from the Philippines. It is the sixth species with six first dorsal spines, a com- pletely serrated preopercular edge, black stomach and intestine, pored lateral line scales from the posttemporal bone to the caudal fin and eight anal fin-rays. The new species is compared with other six-spined (first dorsal) species with these characters and a key is provided. One rare, deep-dwelling Apogon from the Philippines and two undescribed species from the Indian Ocean and West Pacific Ocean are briefly discussed in relation to the new species. Two species have identifiable synonyms: Apogon gularis (Apogon smithvanizi) and Apogon thermalis (Apogon sangiensis). Apogon unicolor is removed from the subgenus Pristicon. A new species of Apogon from deep wa- ter is described from the Philippines. It is the sixth known six-spined species in Apo- gon with a completely serrated preopercular margin, pored lateral-line scales from post- temporal to base of caudal fin, black stom- ach and intestine, and eight anal fin-rays. This description, based on one specimen, is being provided because the unusual depth of the trawl collection makes it unlikely that more material will be available soon. Six spines in the first dorsal fin and a completely serrated preopercular margin are found in relatively few species of the genus Apogon. The pan-tropical subgenus Apo- gon, with about 35 species, has the ventral portion of the preopercle unossified, as a thin fleshly flap, the vertical edge of the preopercle serrated, two supraneural bones, no free uroneurals, ctenoid scales, pored scales forming a complete lateral line, eight anal fin-rays, and the stomach and intestine pale. An apparently related pan-tropical subgenus, Zapogon, with two species, has a free pair of uroneurals, a black pigmented stomach and intestine, and is otherwise similar to the subgenus Apogon. Fraser (1972) included Apogon trimaculatus Cu- vier in Cuvier & Valenciennes, 1828 and Apogon unicolor Déderlein in Jordan & Snyder, 1901 in the subgenus Pristicon based on three supraneural bones, a free pair of uroneurals, strongly serrated infraor- bitals and preopercular ridge. Apogon uni- color is removed from Pristicon based on the presence of a ventral fleshy, unossified flap of the preopercle, small body scales and a black stomach and intestine. This spe- cies may represent another line within Apo- gon. The Atlantic Ocean subgenus Paron- cheilus, with one species, has a partially os- sified ventral preopercular edge, nine anal rays, some canine teeth, cycloid scales, and is otherwise similar to the subgenus Apo- gon. The remaining species of Apogon with six first dorsal spines, have an ossified, ser- rated ventral preopercular edge. These spe- cies have been placed in various subgenera. The subgenera Pristicon (2 species), Yarica (1 species), and a few species of Ostorhin- chus (those with six first dorsal spines at least 3 species: Apogon nigripes, Apogon amboinensis and Apogon lateralis) have pale stomachs and intestines. This small group of Ostorhinchus is under review by VOLUME 111, NUMBER 4 987 Fig. 1. Province, Balayan Bay. me. Other species of Ostorhinchus (at least 107 species) may either have stomachs and intestines pale, speckled or black pigment- ed. The subgenera Brephamia (2-3 spe- cies), and Zoramia (4 species) have a black pigmented stomachs and intestines. Apogon nanus, provisionally allocated by Allen et al. (1994) in Brephamia has pored lateral line scales extending from the posttemporal bone to the caudal fin unlike the other two species. The subgenus Ostorhinchus has species with six (about 10) or seven (at least 107) visible first dorsal spines. Methods Methods of taking and recording meristic data and measurements are given in Fraser & Lachner (1985). All measurements are in millimeters to the nearest 0.1 mm. All pro- portions are based on standard length and all material is reported by standard length rounded to the nearest millimeter, except for the primary type material. All x-ray photographs are in data files maintained by the author. The following acronyms are used to designate institutions and collec- tions cited: BMNH Natural History Muse- um, London; BPBM Bernice P. Bishop Mu- seum, Honolulu; CAS California Academy of Sciences, San Francisco; (SU) Stanford The holotype of Apogon bryx, 42.5 mm standard length, from the Philippines, Luzon Island, Batangas University (collections now housed at CAS); MNHN Museum National d’ Histoire Naturelle, Paris; RMNH Nationaal Natu- urhistorisch Museum, Leiden; RUSI J. L. B. Smith Institute of Ichthyology, Gra- hamstown, South Africa; USNM collec- tions of the former United States National Museum, deposited in the National Muse- um of Natural History, Smithsonian Insti- tution, Washington, D.C. Apogon bryx, new species Fig. 1 Material examined.—Holotype: Apogon bryx CAS 34408; (42.5); Philippines, Lu- zon Island, Batangus Province, Balayan Bay, south of Barrio Nonong Casto, in 146- 155 m. J. E. Norton. 25 Jun 1966. x-rayed. Comparative material.—Apogon atro- gaster Holotype USNM 70249; (46.8); Philippine Is., Western Luzon; x-rayed. Par- atypes USNM 163227; 7(35-48); same data as holotype; x-rayed. Apogon gularis Holotype USNM 225672; (36.1); Red Sea; x-rayed. Apogon kiensis Holotype SU 6514; (56.9); Japan, Wakanoura Kii; x-rayed. Par- atypes SU 6739; 21(47—59) same data as holotype. USNM 71232; 13(30-—60); Japan, Shimizu; x-rayed. Apogon sp., cf. kiensis RUSI 3075; (39-43); Mozambique; x- 988 rayed. RUSI 3074; (31); Mozamibque; x- rayed. RUSI 3073; (44—46); Mozambique, x-rayed. Apogon nanus Paratype USNM 328635; (28); Malaysia, Sabah. Apogon ni- gripes Syntypes BMNH 1864. 11.15.72-74; (37.3), 1865.2.2.23; (42.3); Zanzibar. Apo- gon sangiensis Holotype RMNH 5577; (63.7); Indonesia, Sangi I. Apogon smith- vanizi Paratypes USNM 331174; (46); Bah- rain. BPBM 36421; (49); Bahrain. Apogon thermalis Holotype MNHN 8686; (54.7); Sri Lanka, Trincomalee. Apogon unicolor Holotype USNM 49708; (59.6); x-rayed. Japan, Yokohama. USNM 173833; (85); Australia, NW Cape Arnhem. Apogon sp. USNM 349036; (42); Philippines, Palawan, Puerto Princesa. USNM 260920; (71); Fiji, Lau Group, Matuku Island (19°9'38’S, 179°45'23"E). USNM 260921; 2(49-72); Fiji, Charybdis Reef (17°12'S, 178°0’E). Diagnosis.—An Apogon in the subgenus Ostorhinchus with six first dorsal spines, a completely serrated preopercular edge, black stomach and intestine, eight anal fin- rays, 14-15 pectoral fin-rays, 22 gill rakers (including rudiments) on first arch, no dark stripes or spots on the head, body or fins. Description.—For general body shape see Fig. 1. Proportional measurements (as percentages of standard length, 42.5): body depth 34.6; head length 40.7; eye diameter 10.8; snout length 9.6; bony interorbital width 8.2; upper jaw length 19.8; caudal peduncle depth unknown; caudal peduncle length 27.5; first dorsal-fin spine length 10.6; second dorsal-fin spine length 17.4; third dorsal-fin spine length 15.3; fourth dorsal-fin spine length 12.5; spine in second dorsal fin 12.5; first anal-fin spine length 2.8; second anal-fin spine length 12.0; pec- toral fin length 22.8; pelvic fin length 22.1. Dorsal fin VI-I,9; anal fin II,8; pectoral fin-rays 14 (right), 15 (left); pelvic fin I,5; principal caudal fin-rays 9+8; pored lateral line scales 24; transverse scale rows above lateral line 2; transverse scale rows below lateral line 5; median predorsal scales un- known; circumpeduncular scale rows un- known; gill rakers 22 on first arch, well de- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON veloped gill rakers 21, (1+4 upper, 17+0 lower). Villiform teeth in many rows on the pre- maxilla; one to two rows of villiform teeth on the sides of the dentary; two to three teeth on the palatine and one row on the vomer; none on ectopterygoid, endoptey- goid or basihyal. Supramaxilla absent. Posttemporal smooth to crenulate on posterior margin. Preopercle serrate on vertical and horizontal margins. Infraorbitals smooth. Scales cte- noid on body. Pored lateral-line scales ex- tend from posttemporal to caudal fin. Ver- tebrate 10+ 14. Three supraneural bones, a single supranumerary spine on the first pterygiophore. Three epurals, 5 hypurals and one pair of uroneurals. Anal opening close to the origin of the anal fin. Caudal fin-rays broken. Life colors.—Unknown. Preserved color pattern.—Body, head and fins generally uniform light grayish with no indication of any dark melanophore patterns. Stomach and intestine black, peri- toneum speckled with melanophores. Anal area pale, speckled with a few melano- phores. Distribution.—Known from Batangas Province, Balayan Bay, south of Barrio Nonong Casto, in 146—155 meters by trawl. Etymology.—The Greek noun bryx, in apposition, meaning depth of the sea in ref- erence to the relatively deep water from which this cardinalfish was collected. Remarks.—Apogon bryx has six first dor- sal spines, an uncommon number for nom- inal species of Apogon with a completely serrated preopercle edge, eight anal rays, pored lateral-line scales from the posttem- poral to caudal fin, and a black stomach and/or intestine. Apogon thermalis Cuvier, 1829, Apogon sangiensis Bleeker, 1857, Apogon kiensis Jordan & Snyder, 1901b, a new species related to Apogon kiensis, Apo- gon gularis Fraser & Lachner, 1984, Apo- gon smithvanizi Allen & Randall, 1994, Apogon nanus Allen et al., 1994, and an- VOLUME 111, NUMBER 4 other new species have this combination of characteristics. Apogon thermalis differs from Apogon bryx in having a dark head stripe and bas- icaudal spot and a deeper body depth (36— 39%). The type specimens of Apogon ther- malis from Sri Lanka and Apogon sangien- sis from Indonesia have in common six first dorsal spines, serrated preopercular mar- gins, 14 pectoral fin-rays, smooth posttem- poral edges and similar gill raker counts (2+4-16+0 for A. thermalis and 2+4—- 15+3 for A. sangiensis). The type of Apo- gon thermalis has a dark snout mark con- tinuing behind the eye, ending at the pos- terior edge of the opercle and a small, cen- tered on the lateral line, basicaudal spot. The type of Apogon sangiensis has no re- maining color pattern, however Bleeker (1857) provided a good color description of the type and a follow up color figure in the Atlas (Bleeker, 1873-76: pl. XLI, fig. 4). The color pattern appears very similar to Apogon thermalis. Both types are soft with poor body shapes and the type of Apogon sangiensis lack the premaxilla, so body pro- portions are of limited use for type com- parisons. Based on the above information and examination of other material from the Indian Ocean and West Pacific Ocean, Apo- gon sangiensis is considered a synonym of Apogon thermalis. Apogon kiensis has two stripes on the body, a snout mark, usually 15 pectoral fin- rays and 19—22 (16-18 well developed) gill rakers. The material from Mozambique identified by Smith (1961) as Apogon kien- sis is an undescribed species having a third, shorter, stripe between the two stripes, usu- ally 14 pectoral fin-rays and 22—25 (21-22 well developed) gill rakers. Both species differ from Apogon bryx in preserved color pattern. The undescribed African species has similar pectoral fin-ray and gill raker counts with the new species. Apogon gularis has the anus near the pel- vic fins unlike any other known species of Apogon. In the original description, Apogon smithvanizi was compared with Apogon 989 kiensis but not with Apogon gularis. In ad- dition to having similar preserved color pat- terns (a dark snout mark, a small spots be- hind the eye, generally pale body and fins) and body proportions, Apogon gularis and Apogon smithvanizi have in common: the anus in a forward position near the pelvic fin, black stomach and intestine, usually 14 pectoral fin-rays, six first dorsal spines, a serrated preopercular margin, and 23—26 (20—24 well developed) gill rakers. Both nominal species have been taken only from trawls in deeper waters, 30—40 m for Apo- gon smithvanizi and 60—290 m for Apogon gularis. No differences of substance were found. Apogon smithvanizi is treated here as a synonym of Apogon gularis. Apogon nanus is a small, slender species with more gill-rakers on the first arch (28— 29) than Apogon bryx, a dark spot on the tip of the lower jaw, a narrow dark band along the base of the anal fin and 13 pec- toral fin-rays. Apogon sp. has basicaudal and opercular spots, dark bars or saddles under each dor- sal fin and 15—16 pectoral fin-rays, clearly different from Apogon bryx. Apogon sp. will be described by J. E. Randall and this author as part of their review of Apogon trimaculatus and Apogon rhodopterus. Artificial key for preserved material of Apogon from the Indo-West Pacific region with six first dorsal spines, black stomach and intestine 1. Pored lateral-line scales from posttem- poral to caudal fin 1’. Pored lateral-line scales incomplete, ending before origin of second dorsal fin, remaining scales of lateral line with pits. (Brephamia) (West Pacific Ocean) 2. Dark basicaudal spot present; no stripes on body. (West Pacific Ocean) Werte. 2 Apogon neotes Allen et al., 1994 2'. No dark basicaudal spot; stripes on body. (West Pacific Ocean) Bae ey ae iat, Apogon parvulus . (Smith & Radcliffe in Radcliffe, 1912) 990 10. OF ie Ne We ae . Eight anal fin-rays PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Anal opening near origin of anal fin . . 4 . Anal opening near posterior base of pelvic fins. (Indo-West Pacific Ocean) ... Apogon gularis Fraser & Lachner, 1984 Ventral part of preopercle an unossi- edsinoveable stapes ois eee 5) . Ventral part of preopercle ossified and CdLe SChALC 24 rae coe eee roe Tl Preopercular ridge smooth (Zapogon) 6 . Preopercular ridge serrate. (West Pa- cific Ocean) Apogon unicolor Déderlein in Jordan & Snyder, 1901 Scales above and below lateral-line scales about the same size. (Red Sea) . Apogon isus Randall & Bohlke, 1981 . Scales in rows above lateral-line scales much smaller, more than 35. (Atlantic, Indian and Pacific Oceans) ......... Apogon evermanni Jordan & Snyder, 1904 Nine anal fin-rays Dark stripes on sides of body. (West Paciic Ocean) eee Apogon compressus (Smith & Radcliffe in Radcliffe, 1911) . No dark stripes on sides of body (Zo- ramia). (Indo-West Pacific Ocean)... 9 Dark line on dorsum from origin of first dorsal fin onto caudal peduncle; ventral margin of caudal peduncle with dark line. (Indo-West Pacific Ocean) Apogon leptacanthus Bleeker, 1856 . No dark line on dorsum or on dorsal and ventral surfaces of the caudal pe- duncle Gular area dark; vertical short dark lines just above insertion of some anal rays. (West Bacitic*@cean) 1.2 4-25. Apogon perlitus Fraser & Lachner, 1985 Gular area pale; no vertical lines above andlfiray'sieiie) Salen See ae 11 Opercular flap with prominent to dif- fuse dark spot; caudal spot small with many diffuse melanophores on caudal peduncle. (West Pacific Ocean) .... ... Apogon gilberti (Jordan & Seale, 1905) No spot on opercular flap; caudal spot small, without diffuse melanophores on caudal peduncle. (Indo-West Pacific Ocean) .... Apogon fragilis Smith, 1961 Gill-rakers 19-25; 14-16 pectoral rays 13 Gill-rakers 28—29; 13 pectoral fin rays. (West Pacific Ocean) Apogon nanus Allen et al., 1994 13. Body and/or head with dark marks or stripes . Body and head without dark marks or stripes. (West Pacific Ocean) Apogon bryx, new species 14. Body with dark stripes; no dark basi- Caudal SpOt ss. - choy. <, ere eee 16 14’. No dark body stripes; dark basicaudal SPO te oe os ale 15 15. Dark opercular spot; dark saddle under each dorsal fin; 15-16 pectoral rays. (West Pacific) 22: ae eee Apogon sp. 15’. No dark opercle spot; dark stripe from snout ending on opercle; 14 pectoral rays. (Indo-West Pacific Ocean) .... Apogon thermalis Cuvier in Cuvier & Valenciennes, 1829 16. Two stripes, one over the eye to pos- terior base of second dorsal fin, second from snout to end of caudal fin; 16-18 well developed gill rakers; usually 15 pectoral fin rays. (Indo-West Pacific Ocean) 2... alicia se. eee ... Apogon kiensis Jordan & Snyder, 1901 Three stripes, one over the eye reach- ing on to caudal peduncle, a second from over the eye just above the broader, mid-lateral stripe from snout to tip of caudal fin; 21—22 well devel- oped gill rakers; usually 14 pectoral fin rays. (West Indian Ocean) ...Apogon sp. LIC There is no obvious evidence, in the form of external damage, to suggest that an addi- tional small first dorsal spine was present in the material of Apogon bryx. The x-rays pro- vided no internal evidence of an articulation or support zone for an additional small first dorsal spine (two supranumerary spines in- stead of one). All of the other species with six first dorsal spines discussed here have only one supranumerary spine on the first pterygiophore. The only deep water (from 83 meters), seven-spined species from the Phil- ippines with gill raker counts greater than 20 is Apogon atrogaster (Smith & Radcliffe in Radcliffe 1912) with 26-27 (24 well devel- oped) gillrakers. It has a dark snout mark. Several of the paratypes appear to have six first dorsal spines (Fraser & Lachner 1984). At least one of these paratypes (41.0 mm SL) VOLUME 111, NUMBER 4 has a lower gill raker count (2+5-17+0 = 24) within an expected possible range of var- iation for Apogon bryx. Three other speci- mens are too badly damaged to obtain counts. None of this paratypic material can be iden- tified with any certainty by me. Apogon bryx may be more similar to the undescribed western Indian Ocean species than any of the other species mentioned, based on body shape, pectoral fin-ray and gill raker counts. These two species and Apogon kiensis need to be examined for possible relationships with the Apogon quadrifasicatus complex, all species with seven first dorsal spines. Acknowledgments For the use of museum facilities as a part of my long-term studies of cardinalfishes, I extend thanks to P. J. RP Whitehead (BMNH), A. Y. Suzumoto and J. E. Randall (BPBM), D. Catania, W. N. Eschmeyer and T. Iwa- moto (CAS), M. Boeseman (RMNH), M. M. Smith (RUSD, and M. L. Bauchot (MNHN). S. L. Jewett, D. G. Smith, J. T. Williams of the Smithsonian Institution (USNM), all aid- ed in curatorial processes and during my vis- its. D. G. Smith and J. T. Williams helped with photography and x-rays. Leonard P. Schultz funds and office space were provid- ed by V. G. Springer (USNM) for several study trips to the Smithsonian. J. E. Randall and G. R. Allen (WAM) and an anonymous reviewer provided useful comments for drafts of the manuscript. Literature Cited Allen, G. R., & J. E. Randall. 1994. A new species of cardinalfish (Apogon: Apogonidae) from Ara- bian Seas.—Revue frangaise d’ Aquariologie 21(1994)1—2:24—26. , R. H. Kuiter, & J. E. Randall. 1994. Descrip- tions of five new species of cardinalfishes (Apo- gonidae: Apogon) from Maumere Bay, Flores, Indonesia and surrounding regions.—Revue frangaise d’ Aquariologie 21(1994)1—2:27-38. Bleeker, P. 1856. Zevende bijdrage tot de kennis der ichthyologische fauna van Ternate (1).—Natu- urkundig Tijdschrift voor Nederlandsch-Indié 10:357—386. 991 . 1857. Bijdrage tot kennis der ichthyologische faun van de Sangi-eilanden.—Natuurkundig Tijdschrift voor Nederlandsch-Indié 13:369— 380. . 1873-1876. Atlas ichthyologique des Indes Orientales Néédandaises. Fréderic Muller, Am- sterdam, 7:126 pp., 42 col. pls. Cuvier, G., & A. Valenciennes. 1828. Histoire Natu- relle des Poissons. Paris (De Luxe ed), 2:1—371. , . 1829. Histoire Naturelle des Pois- sons. Paris (De Luxe ed.), 3:1—368. Fraser, T. H. 1972. Comparative osteology of the shal- low water cardinal fishes (Perciformes: Apo- gonidae) with reference to the systematics and evolution of the family.—Ichthyological Bulle- tin (34):1—105. , & E. A. Lachner. 1984. An unusual Indo-West Pacific cardinalfish of the genus Apogon (Te- leoste1: Apogonidae).—Proceedings of the Bi- ological Society of Washington 97(3):632—636, 2 figs. , & . 1985. A revision of the cardinal- fish subgenera Pristiapogon and Zoramia of the Indo-Pacific region (Teleostei: Apogonidae).— Smithsonian Contributions to Zoology 412:1— 47. Jordan, D. S. & A. Seale. 1906. The fishes of Samoa. Description of the species found in the Archi- pelago with a provisional check list of the fishes of Oceania.—Bulletin of the Bureau of Fisher- ies 25:175—488. , & J. O. Snyder. 190 1a. List of fishes collected in 1883 and 1885 by Pierre Louis Jouy and pre- served in the United States National Museum with descriptions of six new species.—Proceed- ings of the United States National Museum 23(1235):739-769. , & . 1901b. A review of the cardinal fishes of Japan.—Proceedings of the United States National Museum 23(1204):891—913. Radcliffe, L. 1911. Notes on some fishes of the genus Amia, family of Cheilodipteridae, with descrip- tions of four new species from the Philippine Islands.—Proceedings of the United States Na- tional Museum 41(1853):245—261. . 1912. Descriptions of fifteen new fishes of the family Cheilodipteridae from the Philippine Is- lands and contiguous waters.—Proceedings of the United States National Museum 41(1868): 431-446. Randall, J. E., & J. E. Bohlke. 1981. Status of the cardinalfishes Apogon evermanni and A. aniso- lepis (Perciformes: Apogonidae) with a descrip- tion of a related new species from the Red Sea.—Proceeding of the Academy of Natural Sciences of Philadelphia 133:129—140. Smith, J. L. B. 1961. Fishes of the family Apogonidae of the western Indian Ocean and Red Sea.— Ichthyological Bulletin 22:373—418. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):992-1007. 1998. The Neotropical catfish genus Epapterus Cope (Siluriformes: Auchenipteridae): a reappraisal Richard P. Vari and Carl J. Ferraris, Jr. (RPV) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.; (CJF) Department of Ichthyology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. Abstract.—The Neotropical auchenipterid catfish genus Epapterus Cope is reviewed. Epapterus was found to share synapomorphies with Auchenipterus Valenciennes and Pseudepapterus Steindachner and within that clade it is the sister-group to Pseudepapterus. Epapterus lacks an adipose dorsal fin, a derived reductive feature unique to the genus within the clade consisting of Epapterus, Auchenipterus, and Pseudepapterus and which is hypothesized to be a syna- pomorphy for the genus. The extension of the interradial membrane to join the contralateral pelvic fins is identified as an additional possible synapomorphy for Epapterus. Contrary to recent practice, Epapterus is considered to consist of only two species, E. dispilurus Cope (1878) and E. blohmi Vari et al. (1984). Examination of Epapterus samples from a number of localities in the Rio Amazonas and Rio Paraguay basins failed to reveal differences between the populations of the genus in those drainage systems. As a consequence Epap- terus chaquensis Risso & Risso (1962), described from the Rio Paraguay sys- tem, is placed as a synonym of E. dispilurus originally described from the western portion of the Amazon basin. Euanemus longipinnis Steindachner (1881) is retained as a synonym of Epapterus dispilurus as proposed by Stein- dachner (1882) and Eigenmann & Eigenmann (1888). Epapterus dispilurus is found to have a broad distribution in the central and western portions of the Rio Amazonas system and the Rio Paraguay basin. The distribution of E. bloh- mi is extended from the state of Guarico, Venezuela to include other portions of the Rio Orinoco basin and the Rio Tuy of the Caribbean coast of Venezuela. Resumo.—O género de bagres neotropicais Epapterus Cope € revisado. Epapterus compartilha varias sinapomorfias com Auchenipterus Valenciennes e Pseudepapterus Steindachner, e dentro deste grupo é 0 grupo irmao de Pseu- depapterus. Epapterus nao possui nadadeira adiposa, uma caracteristica redu- tiva derivada Unica para o género dentro do grupo composto por Epapterus, Auchenipterus e Pseudepapterus, e proposta como sinapomorfica para 0 gé- nero. A extenséo da membrana inter-radial que une as duas nadadeiras pélvicas é uma outra possivel sinapomorfia para Epapterus. Contrariamente a opiniao corrente, Epapterus € composto de apenas duas espécies, E. dispilurus Cope (1878) e E. blohmi Vari et al. (1984). O exame de amostras de Epapterus provenientes de varias localidades no rio Amazonas e Rio Paraguay nao revelou nenhuma diferenga entre as populagdes daquelas duas drenagens. Consequen- temente, E. chaquensis Risso & Risso (1962), descrito da bacia do Rio Para- guay, é colocado como sinénimo de E. dispilurus, originalmente descrito da porgao ocidental da bacia amazénica. Euanemus longipinnis Steindachner (1881) é mantido como sinénimo de E. dispilurus conforme proposto por Stein- dachner (1882) e Eigenmann & Eigenmann (1888). Epapterus dispilurus possui VOLUME 111, NUMBER 4 993 ampla distribui¢ao nas por¢oes central e ocidental do sistema do rio Amazonas, assim como na bacia do Rio Paraguay. A distribuigao geografica de E. blohmi é ampliada do estado de Guarico, Venezuela, para outras porgdes da bacia do rio Orinoco e ao norte para o Rio Tuy na costa caribenha de Venezuela. In his original description of the auchen- ipterid catfish genus Epapterus from the Pe- ruvian Amazon, Cope (1878:677) distin- guished the genus and its single species from the other then-known genera now as- signed to the Auchenipteridae by a combi- nation of characters, most notably the lack of the adipose fin, the reduced dorsal fin, and the absence of jaw teeth. Soon there- after Steindachner (1881) described another auchenipterid, Euanemus longipinnis, from specimens collected in the western Amazon along the present Peru-Brazil border. Stein- dachner (1882:31) noted that the two nom- inal species were apparently identical and Eigenmann & Eigenmann (1888:152) for- mally proposed that Euanemus longipinnis was a synonym of Epapterus dispilurus. Soon thereafter Eigenmann & Eigenmann (1890) redescribed E. dispilurus on the ba- sis of specimens that they considered to be syntypes of Euanemus longipinnis (see, however, under ““Remarks”’ for Epapterus dispilurus). The seven decades following this series of publications saw only one ci- tation of Epapterus based on additional ma- terial, that of Fowler (1940) of a single specimen of FE. dispilurus collected in the Rio Ucayali in Amazonian Peru. This hiatus ended when Risso & Risso (1962) de- scribed E. chaquensis from five specimens collected in the Rio Paraguay basin of Ar- gentina. Risso & Risso distinguished their nominal species from E. dispilurus on the basis of purported differences in the num- ber of pectoral- and anal-fin rays, relative eye size, and details of body and fin pig- mentation. In the absence of available sam- ples of E. dispilurus, Risso & Risso com- pared E. chaquensis to data in the original description of E. dispilurus by Cope (1878) and a relatively simple drawing of the lec- totype of the latter species published twice by Fowler (1941:468, fig. 26; 1945:66, fig. 26). Vari et al. (1984) described another spe- cies, E. blohmi, from samples originating in the central portions of the Rio Orinoco sys- tem. Epapterus blohmi is readily differen- tiated from nominal congeners by its dis- tinct caudal-fin pigmentation pattern, a dif- ference supplemented by other features. In their discussion of E. blohmi Vari et al. (1984) also commented on the pronounced sexual dimorphism in the species of Epap- terus and noted a series of characters po- tentially informative as to the phylogenetic position of the genus within the Auchen- ipteridae. Those authors noted, however, that definitive statements on the questions of the monophyly and relationships of Epapterus necessitated an encompassing phylogenetic analysis of the Auchenipteri- dae, an effort which was beyond the scope of their study. Vari et al. (1984) also briefly reported on new locality records for the two other Epapterus species (E. dispilurus and E. chaquensis) generally recognized as val- id at that time, but lacked the population samples necessary to address the utility of the purported distinguishing features of those species. In the course of our ongoing studies within the Auchenipterinae (Ferraris & Vari 1999) we have been able to examine nu- merous specimens of Epapterus in addition to those available to previous authors. This additional material led to a reappraisal of the distinctiveness of E. dispilurus and E. chaquensis. We address that problem here- in, discuss the phylogenetic information pertinent to the monophyly of Epapterus, note broader meristic variation within E£. blohmi revealed by the examination of ad- 994 ditional specimens, and report extensions of the known distribution of E. blohmi. Materials and Methods Materials are listed alphabetically by country and within country by district, state, or province, and then repository ab- breviation. Specimen sizes are reported in standard length (SL). Localities of speci- mens, other than type-localities, are trans- lated into English when listed originally in another language. Coordinates are either taken from labels associated with speci- mens, or from publications, information provided by collectors, or from gazetteers when the locality is sufficiently precise. Institutional abbreviations used are as follows: ANSP, Academy of Natural Sci- ences of Philadelphia; MCNG, Museu de Ciencias Naturales, Guanare; MCZ, Muse- um of Comparative Zoology, Cambridge; MHNG, Muséum d’ Histoire naturelle, Ge- neva; MLP, Museo de La Plata; MUSM, Museo de Historia Natural de la Universi- dad Nacional Mayor de San Marcos, Lima; MZUSP, Museu de Zoologia, Universidade de Sao Paulo; NMW, Naturhistorisches Mu- seum Wien, Vienna; and USNM, National Museum of Natural History, Smithsonian Institution, Washington, D.C. Vertebrae and unpaired fin rays were counted, when possible, from radiographs. Comparative counts made directly from specimens often failed to discern the ante- rior most anal-fin ray(s) and the posterior most dorsal- and anal-fin rays in small spec- imens. Vertebral counts were separated into precaudal and caudal elements. The anterior most caudal vertebra is the element pos- sessing an elongate hemal spine that ex- tended just anterior to the first basal radial of the anal fin. Usually, the last precaudal vertebra possesses a markedly shorter he- mal spine than does the first caudal verte- bra, and is the anterior most vertebra with a closed hemal arch. The ural complex is counted as one element. Precaudal and total vertebral counts assume that the Weberian PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON complex is composed of four centra. The fifth centrum is suturally united to the com- plex and lacks articulated ribs. The first rib- bearing centrum is the sixth vertebra. Fin-ray counts include all elements and treat the two posterior most dorsal- and anal-fin rays that articulate on the last distal radial as separate rays. The last pelvic-fin ray is often branched at its base, which is hidden beneath thick skin. Dissection was sometimes performed to verify the pelvic- fin ray count. Caudal-fin ray counts include only the branched principal rays. Anal-fin ray counts from sexually mature, trans- formed males assume that the gonopodium consists of one unbranched and one branched ray in the few instances in which separate elements could not be discerned from radiographs. The posterior most anal- fin rays are recorded as branched, although many were too small and/or fine to discern whether they actually branch. Rudimentary anal-fin rays occur irregularly. Rudiments that appear to possess a base that articulates with a basal radial element were included in counts, even if the rays do not extend distally to the fin margin. The first two rays posterior to the gonopodium in males often exhibit this rudimentary development. Ru- diments suspended in the fin margin and fully formed rays that do not appear to ar- ticulate with a radial element were not counted. Gill rakers were counted on the first gill bar on the right side of the specimen. Counts included all rakers, including ante- rior rudiments. Measurements were made with digital calipers and were point-to-point, as follows: body depth was taken at the anal-fin origin; head length was measured parallel to the body axis, from the posterior tip of the bony operculum to the snout tip; length of anal- fin base was measured from the posterior edge of the anus to the insertion of the last anal-fin ray; orbital width was taken as the horizontal measure across the middle of the eye, bounded by the infraorbital bones; pel- vic-fin length represents the distance from VOLUME 111, NUMBER 4 the attachment of the first branched ray to the fin tip; pectoral-fin spine length is from the anterior (or lateral) spine base to the tip of the bony spine not including the fleshy or flexible bony terminal parts; snout length is from the snout tip to the anterior margin of the eye. Proportional measurements are presented as percentages of standard length (SL) or head length (HL). Values for the lectotype of Epapterus dispilurus are indicated in brackets in the description. Nuptial males are those individuals dem- onstrating the transitory sexually dimorphic features of the maxillary barbels and dorsal- fin spine. Epapterus Cope Epapterus Cope, 1878:677 (type species: Epapterus dispilurus Cope, 1878, by monotypy).—Eigenmann & Eigenmann, 1888:152 [citation]—Eigenmann & Ei- genmann, 1890:292 [citation].—Eigen- mann, 1910:396 [citation].—Fowler 1951:465 [citation].—Vari et al., 1984: 468 [discussion of relationships and po- tential synapomorphies]. Diagnosis.—Epapaterus along with Au- chenipterus Valenciennes, in Cuvier and Valenciennes (1840) and Pseudepapterus Steindachner (1915) form a monophyletic assemblage within the Auchenipteridae de- fined by various synapomorphies (Ferraris 1988, Ferraris & Vari 1999). The combi- nation of the absence of jaw teeth and the possession of a reduced dorsal fin with only two to four short rays and a poorly devel- oped spine (other than in nuptial males) de- limits a clade formed by Epapterus and Pseudepapterus within the Auchenipteri- dae. The lack of an adipose fin is a syna- pomorphy for the species of Epapterus within that lineage. Epapterus species also have the pelvic fins joined to each other across the ventral midline by extensions of the interradial membrane; an additional po- tential synapomorphy for the members of the genus (see comments under “‘Phyloge- 995 netic relationships and monophyly of Epap- terus’ below). Phylogenetic relationships and mono- phyly of Epapterus.—In his original de- scription of Epapterus, Cope (1878:677) listed a series of “‘Char. Gen.”’, presumably the diagnostic features for the genus. These features largely consisted of what would now be regarded as various plesiomorphic characters, at least at the level of the Au- chenipteridae. Nonetheless, three features cited by Cope as distinguishing Epapterus from Euanemus (=Auchenipterus), the lack of an adipose fin, the reduced number of dorsal-fin rays, and the absence of teeth on the jaw and palate, are considered derived within the Auchenipteridae under the con- text of an encompassing phylogenetic anal- ysis (Ferraris 1988). Steindachner (1915) subsequently described Pseudepapterus as a subgenus of Auchenipterus. Pseudepapa- terus shares with Epapterus a lack of jaw dentition and a reduced dorsal fin but was distinguished from the latter genus on the basis of the possession of an adipose fin. Mees (1974:112) commented briefly on the similarities between Auchenipterus, Epa- paterus, and Pseudepapterus. Vari et al. (1984:467—468) identified a number of fea- tures in Epapterus of potential phylogenetic importance. Those authors were unable to make definitive statements on these ques- tions in the absence of comparative speci- mens of Pseudepapterus, and as a conse- quence of the lack, at that time, of a rig- orous hypotheses of higher level groupings within the Auchenipteridae. Auchenipterus, Epapterus, and Pseude- papterus comprise a monophyletic group within the Auchenipteridae united by vari- ous characters summarized by Ferraris (1988) and Ferraris & Vari (1999). The re- duced dorsal fin with only two to four short rays and a poorly-developed spine other than in nuptial males, along with the lack of dentition on both the jaws and palate, are derived features within the Auchenipteridae uniting Pseudepapterus and Epapterus as sister-taxa. Epapterus has been traditionally 996 distinguished by its lack of an adipose dor- sal fin. An adipose fin is also absent else- where in the Auchenipteridae in Trachel- yopterus Valenciennes, Trachelyopteri- chthys Bleeker, and Trachelyichthys Mees (the reported absence of the fin in Cerato- cheilus by Miranda-Ribeiro (1918) was in error; see comments in Ferraris & Vari (1999). The lack of an adipose fin is con- sidered synapomorphic for the species of Epapterus in the clade consisting of Au- chenipterus, Epapterus, and Pseudepapte- rus under the overall most parsimonious hypothesis of intrafamilial relationships for the Auchenipteridae (Ferraris 1988). Epapterus species also have the pelvic fins joined to each other across the ventral midline by extensions of the interradial membrane. Such continuity of the contra- lateral pelvic fins is hypothesized as derived within the Auchenipteridae given the ab- sence of a comparable union of the fins in most auchenipterids and in proximate out- groups to the family. A continuity between the contralateral pelvic fins also occurs else- where within the clade consisting of Au- chenipterus, Epapterus, and Pseudepapte- rus in Auchenipterus fordicei Eigenmann & Eigenmann (Ferraris & Vari 1999). Auchen- ipterus fordicei, a poorly known species, shares with its congeners the presence of grooves on the ventral surface of the head that accommodate adducted mental barbels, a synapomorphy for the species of Auchen- ipterus (Ferraris & Vari 1999). Nuptial males of A. fordicei are unknown and it is uncertain whether A. fordicei has the sec- ond potential synapomorphy for Auchenip- terus, the presence of papillae on the dorsal and medial surfaces of the ossified maxil- lary barbel of nuptial males (Ferraris & Vari 1999). Auchenipterus fordicei, nonetheless, lacks the derived features considered syna- pomorphic for Epapterus and Pseudepap- terus and retains a relatively large dorsal fin and an adipose fin, contrary to the derived reduction and absence, respectively, of those structures in Epapterus. Thus, the common possession of pelvic fins joined to PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON each other across the ventral midline by ex- tensions of the interradial membrane in Au- chenipterus fordicei and Epapterus is most parsimoniously considered a result of con- vergence. Key to the species of Epapterus 1. Dorsal lobe of caudal fin with irregularly rounded patch of dark pigmentation in central portion of lobe, pigmentation not forming distinct transverse bar; ventral lobe of caudal fin with distinct patch of dark pigmentation in middle of lobe; pectoral-fin rays 11 to 13, typically 12 or 13 y's dotwea ee Wane ee E. dispilurus — Dorsal lobe of caudal fin with a dark transverse bar extending from upper an- terodorsal margin to posterior margin of middle rays of dorsal lobe; ventral lobe of caudal fin without a distinct patch of dark pigmentation; pectoral-fin rays 9 to Me Sascha. os Ree eee E. blohmi Epapterus dispilurus Cope, 1878 Figs. 1-3, Table 1 Epapterus dispilurus Cope, 1878:677 (type locality: Peruvian Amazon).—Steindach- ner, 1882:31 [Euanemus longipinnis Steindachner, 1881:17 equated with Epapterus dispilurus|.—Eigenmann & Eigenmann, 1888:152 [citation].—Eigen- mann & Eigenmann, 1890:292 [rede- scription based on specimens erroneously thought to be syntypes of Euanemus lon- gipinnis|—Eigenmann, 1910:397 [cita- tion].—Fowler, 1915:222 [citation; orig- inal Cope specimens cited as cotypes].— Fowler, 1940:231 [Peru, Rio Ucayali].— Fowler, 1941:468, fig. 26 [lectotype des- ignation].—Fowler, 1945:66, fig. 26 [re- production of Fowler, 1941].—Fowler, 1951:465 [citation].—B6hlke, 1984:24 [type holdings at ANSP].—Vari et al., 1984:470, fig. 2 [Brazil, Ihla da Mar- chantaria; Peru, Rio Ucayali basin; com- parison with Epapterus blohmi]|.—Ortega & Vari, 1986:14 [as component of Peru- vian freshwater fish fauna]. VOLUME 111, NUMBER 4 997 Fig. 1. of dorsal-fin spine and maxillary barbel. Euanemus longipinnis Steindachner, 1881: 17 (type locality: Hyavary [=Brazil, Amazonas, Rio Javari]).—Steindachner, 1882:31 [equated with Epapterus dispi- lurus|.—Eigenmann & Eigenmann, 1888:152 [citation, as synonym of Epap- terus dispilurus|.—Eigenmann & Eigen- mann, 1890:292 [citation, as synonym of Epapterus dispilurus|.—Eigenmann, 1910:397 [citation, as synonym of Epap- terus dispilurus|.—Fowler, 1951:466 [ci- tation, as synonym of Epapterus dispi- lurus |. Epapterus chaquensis Risso & Risso, 1962: 5, figs. 1-3 (type locality: Argentina, Chaco, Resistencia).—Risso & Risso, 1964:5 [citation].—Vari et al., 1984:470, fig. 3 [Paraguay, Rio Negro of Rio Par- aguay].—[not Burgess, 1989, pl. 113, un- numbered figure]. Auchenipterus nuchalis, Sands, 1984:24, unnumbered photo.—Burgess, 1989; pl. ie Auchenipterus demerarae, Sands, 1986:43, unnumbered photos. Diagnosis.—Epapterus dispilurus is di- agnosed by, and is readily distinguished from its single congener, E. blohmi, by hav- ing a distinct, irregularly-shaped, patch of dark pigmentation on each caudal-fin lobe which contrasts with the dark transverse bar extending from the upper anterodorsal mar- gin of the dorsal lobe of the caudal fin to the posterior margin of the middle rays of Epapterus dispilurus, nuptial male, USNM 273591, 106 mm SL; showing pronounced development the dorsal lobe, and by the lack of a distinct patch of dark pigmentation on the ventral lobe of the caudal fin in E. blohmi. The two species also differ in the number of pecto- ral-fin rays (11 to 13, typically 12 or 13, in E. dispilurus versus 9 to 11 in E. blohmi), and in differences, other than for nuptial males of E. dispilurus, in the relative length of the maxillary barbel (166—194% of HL in E. blohmi versus 122—163% of HL in E. dispilurus). Epapterus dispilurus nuptial males also have a highly developed dorsal spine with well developed basal and distal anterior projections, modifications which are unknown in E. blohmi. Description.—Body elongate, distinctly compressed, head depressed (Figs. 1, 2). Dorsal profile of head flat or barely convex. Dorsal profile of body from rear of head to end of caudal peduncle slightly to distinctly convex. Ventral profile of head slightly con- vex. Ventral profile of body gently curved to anal-fin origin. Intromittent organ of males extending along entire anterior mar- gin of third anal-fin ray. Anal-fin base gent- ly convex to caudal peduncle. Greatest body depth 19.3—25.6 [19.6] of SL; snout tip to dorsal-fin origin 19.3—22.5 [19.7] of SL; snout tip to pelvic-fin origin 33.2—37.1 [34.0] of SL; snout tip to anal- fin origin 39.0—45.6 [39.6] of SL. Head depressed. Snout viewed from dor- sal view somewhat more truncate in mature males than in females in which snout mar- 998 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. maxillary barbel in females and immature individuals. gin distinctly rounded from that view. Head length 18.3—21.6 [18.9] of SL. Lower jaw slightly shorter than upper jaw in females and immature specimens, difference in jaw lengths slightly more pronounced in mature males; no teeth present on premaxillae, den- taries, vomer, and palatines. Lower pharyn- geal tooth plates large and round with short conical teeth; convex fourth upper pharyn- geal tooth plate with short conical teeth. Snout length 36.2—40.9 [40.7] of HL. Or- bital margin not free, horizontal width of orbit 26.7—30.2 [29.2] of HL. Length of postorbital portion of head 41.5—46.7 [41.5] of HL. Interorbital region gently convex. Nares of each side of head separated by dis- tance approximately equal to 4.5—5.0 times diameter of posterior nostril; anterior nostril somewhat tubular, located on anterodorsal surface of snout, above lip; posterior nostril larger, oval. Gill rakers on first arch 33 to 42. Bran- chiostegal rays 6. Gill membranes broadly attached to isthmus. Maxillary barbels rounded in cross section, elongate, length 122-163% of HL in females, immature specimens, and males without fully devel- oped ossification of barbel; barbel length 191% of HL in single available nuptial male; tip of barbel reaching posteriorly to, nearly or to, end of pectoral fin in females, immature specimens, and males without fully developed ossification of the barbel, extending distinctly beyond tip of pectoral fin in fully nuptial male; degree of ossifi- Epapterus dispilurus, female, USNM 305651, 102 mm SL; showing form of dorsal-fin spine and cation of barbel highly sexually dimorphic (see “Sexual dimorphism” below). Mental barbels four, transversely rounded, arranged in arc along ventral surface of jaw, barbels reaching posteriorly approximately to line through middle of pectoral fin. Lateral line complete, dendritic, with short side branches at irregular intervals, di- vided on caudal fin into 2 or 3 branches that run in some individuals through slightly os- sified tubes. Dorsal fin greatly reduced, with 1 spine and 2 or 3 rays (2 rays present in 1 of 43 specimens examined for this feature). Dor- sal-fin spine non-serrate and sexually di- morphic, relatively short and slender, length 2.8-3.6 of SL in females, immatures, and males not in reproductive condition. Dorsal fin proportionally much thicker and dis- tinctly elongate in one available nuptial male, length 16.4 of SL (see “Sexual di- morphism’”’ below for detailed description of spine). First dorsal-fin ray typically slightly longer than dorsal-fin spine in fe- males and immature specimens. Single available nuptial male with first dorsal-fin ray distinctly longer proportional to SL than in females, immature specimens, and males not in reproductive condition, but with lengthened ray extending only about three- quarters of length of greatly enlarged dor- sal-fin spine. Adipose dorsal fin absent. Pectoral fin with 1 spine and 11 to 13, most often 12 [12], rays; pectoral-spine length 12.3—14.0 [13.0] of SL; medial mar- VOLUME 111, NUMBER 4 gin of pectoral spine with series of serra- tions, serrations absent basally and some- times along distal one-fifth of spine; rayed portion of fin pointed, longest pectoral-fin rays ranging from approximately same length as, to somewhat longer than, pectoral spine; length of longest pectoral-fin rays 12.3-14.4 of SL; tip of pectoral fin not reaching to pelvic-fin origin. Pelvic-fin margin pointed, fin with 14 to 16 rays [15], lateral most rays longest, length of longest pelvic-fin rays 16.0—19.3 of SL; pelvic fin with broad, posteroven- trally sloping base with interradial mem- brane continuing across midline of body to join interradial membrane of contralateral pelvic fin; insertion of pelvic fin situated distinctly dorsal of ventral margin of body. Anal-fin margin in females smoothly convex anteriorly, straight for much of its length and convex posteriorly; last un- branched and first branched anal-fin rays in males greatly developed and conjoined for support of intromittent organ (see “‘Sexual dimorphism” below for details); remainder of anal fin of males as in females; length of anal-fin base 51.4—56.0 [55.5] of SL. Anal- fin rays 54 to 61 [59]. Caudal fin distinctly emarginate. Precaudal vertebrae 14 to 16, typically 15 [15], caudal vertebrae 33 to 36 [35], total vertebrae 48 to 50 [50]. Color in life.-—Sands (1986:43, unnum- bered photos) published two photographs of what is apparently the same individual of Epapterus dispilurus (identified by that au- thor as Auchenipterus demerarae). These il- lustrations were later reproduced by Bur- gess (1989, pl. 113, unnumbered photo- graphs) who identified the species as Au- chenipterus nuchalis. Overall dark pigmentation on the head and body in spec- imen in the photos is as described below for preserved specimens, but with more lightly pigmented portions of body having a silvery sheen which also extends onto the basal portions of the caudal and anal fins. The anterior margin of the pectoral fin white. Barbels with white tips. Neither the 999 silvery pigmentation nor the white margin to the pectoral fin is apparent in the aquar- ium specimen of E. dispilurus published by Sands (1984:24, photo; identified therein as Auchenipterus nuchalis). Color in alcohol.—Overall ground col- oration ranging from tan to light purplish brown, overall pigmentation tending to in- crease with increasing body size, but with population samples of similar size individ- uals from different sites in the Rio Ucayali basin showing notable difference in overall intensity of pigmentation. Dorsal surface of head in lighter colored specimens with scattered chromatophores more concentrated lateral to, and particu- larly posterior to, fronto-parietal fontanel. Margin of upper lip in region anterior to orbit with distinct patch of dark pigmenta- tion. With increasing intensity of overall head and body pigmentation, chromato- phore field on dorsal surface of head ex- pands laterally and anteriorly to contact or- bit laterally and patch of pigmentation on upper lip anteriorly. In very dark individu- als entire dorsal portion of head dark, other than for light anterior margin of upper lip. Lateral surface of head with scattered dark chromatophores in light specimens; darker specimens with patches of dark pigmenta- tion anterior and posterior to orbit contin- uous dorsally with dark pigmentation on dorsal portion of head. Dorsal portion of body with scattered dark chromatophores in lighter specimens, but with nearly solid purplish pigmentation in darker individuals. Distinct, dark humer- al spot present in all individuals; spot slightly horizontally elongate in some smaller, lighter-colored specimens, more rounded in larger specimens, with posterior portion of spot continuous in some individ- uals with variably developed zigzag pattern of dark chromatophores overlying lateral line. Darker individuals with ventral and sometimes midlateral portions of body with dusky purplish pigmentation. Caudal fin with irregular patch of dark pigmentation on dorsal lobe; vertical extent 1000 of patch usually greater than its horizontal length. Ventral lobe of caudal fin with ir- regular patch of dark pigmentation on cen- ter of lobe; shape of patch ranging from somewhat rotund to vertically elongate. Caudal pigmentation more intense in spec- imens with overall darker coloration. Anal fin nearly unpigmented in some specimens, with distal margin darker in most examined specimens. Dark pigmentation on distal portions of fin limited to anterior one-half to two-thirds of fin in some specimens whereas other individuals have field of pig- mentation along distal region of fin more elongate, occasionally with fin margin quite dark along its entire length. Basal portions of anal fin purplish in very dark individuals. Extent of dark pelvic fin pigmentation var- iable among specimens with differences in degree of development correlated with overall intensity of dark pigmentation of specimen. Distal margin of pelvic fin dusky in all specimens, more so in dark individ- uals. Lateral margin and basal portions of pelvic fin in less intensely pigmented indi- viduals with scattered dark chromatophores. Pelvic fin with dark patch of pigmentation in more intensely pigmented specimens. Pectoral fin with scattered dark chromato- phores more concentrated along distal mar- gin; more intensely pigmented individuals with varyingly developed basal patch of dark pigmentation. Sexual dimorphism.—Females attain a greater maximum size than do males. The largest specimen examined by us was a 124 mm female and a number of females ex- ceeded 110 mm. The largest identifiable male was 107 mm and the smallest was 97 mm. The snout in nuptial males is some- what more truncate when viewed from a dorsal view than that in females in which the snout margin is distinctly rounded. Fe- males have the bony core of the maxillary barbel relatively short with the remainder of the barbel flexible, thin, and extending pos- teriorly to the middle of, or to the posterior margin of, the pectoral fin. In the single ex- amined nuptial male the barbel is greatly PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON thickened, distinctly curved (Fig. 1), and somewhat lengthened relative to the con- dition in females. When adducted the barbel of the nuptial male reaches distinctly be- yond the posterior tip of the pectoral fin. The dorsal-fin spine in the single avail- able nuptial male is distinctly thicker and more notably is distinctly longer than the spines of immature individuals, females, and males not in reproductive condition (compare Figs. 1 and 2). The spine in nup- tial males is somewhat sinusoidal in lateral view, with a distinct anterior process to- ward its base and a well developed anterior process toward its tip which gives the distal portion of the spine a harpoon-like shape in lateral view (Fig. 1). The first dorsal-fin ray is also distinctly longer in the nuptial male than in immature specimens, females, and males not in reproductive condition. The dorsal-fin spine of males can be hyperex- tended anteriorly to an approximately 75° angle relative to the predorsal profile, in- stead of the vertical or slightly posterodor- sal fully adducted position of the spine in females and juveniles. Mature males have an anal fin with the last unbranched and the first branched anal-fin rays distinctly thick- ened and more elongate than those in fe- males and conjoined to form the structural support of an intromittent organ. The gen- ital pore of males is at the end of an elon- gate tube that extends along, and is bound to, the anterior margin of the anal fin. The tube terminates slightly beyond the end of the tip of the conjoined last unbranched and first branched anal-fin rays. Distribution.—Central and western por- tions of the Amazon basin along and south of the main channel of the Rio Amazonas, and the Rio Paraguay system in Paraguay, northern Argentina, and southern Brazil (Eie3): Remarks.—Epapterus dispilurus Cope (1878) was described on the basis of two specimens which originated in the upper Amazon basin of Peru. These specimens were cited as “‘cotypes”’ by Fowler (1915: 222). Some years later, and without any dis- VOLUME 111, NUMBER 4 ieee 1001 PHN \ \ 1 d = Sie oF SX —e od Sy ss oN a= ot we LG igi gl y l : i Ly ZN = ; w!, on ee \ ae Aye Va f nN a C7 ( € ae zi eee ay \ ii SE ad) & HIRES SSS: lass ec : ou ? ‘ ) ( Le) DNS 2 V2 \ ( XJ SSCs Sane Fig. 3. Map of central portions of South America showing known distribution of Epapterus dispilurus (type locality of E. dispilurus inexact = Peruvian Amazon; 1 type locality of Euanemus longipinnis; 2 = type locality of Epapterus chaquensis; some symbols represent more than one locality or lot of specimens). cussion, that author (Fowler 1941:468, fig. 26 repeated in Fowler 1945:66, fig. 26) pro- vided a cryptic, but valid, lectotype desig- nation by the statement “tipo, largo 125 mm” which accompanied his line drawing of the species. The cited length corresponds with the total length of the larger of the two syntypes of E. dispilurus which we conse- quently consider to be the lectotype (ANSP 21353). This obscure lectotype designation was overlooked in Bohlke (1984). The name Euanemus longipinnis is gen- erally cited as being published in 1882 with the author credited as being Agassiz in a Steindachner paper. Both this date and au- thor are questionable. The name was pub- lished as the first article of the Denk- schntten der Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse for 1882, but Troschel (1882) listed the paper as having been published in 1881. Barbara Hertzig (NMW, in litt.) indicated that separates of the publication were avail- able, and probably distributed, in 1881, which would account for the date cited in Troschel’s account. On the basis of this ev- idence, we use the year 1881 as the date of description of E. longipinnis. Steindachner (1881:17) indicated that the specimens on which he based his descrip- tion of the species were sent to him by Ag- assiz with the name “‘Euanemus longipinnis Agass.”’ Agassiz’s contribution appears to be limited to the creation of the name and sending the specimens to Steindachner giv- en that the format of the species description 1002 is typical for Steindachner’s other publica- tions of the period. As such, we consider Steindachner as the author of the name and only the specimens examined by Steindach- ner can be considered as constituting the type series. Steindachner (1881:18) noted in the original description of the species that he examined four specimens, a number matching the identified syntypes in the NMW holdings (NMW 46682:1—4). We have examined two of the four syntypes catalogued as NMW 46682:1-—4 and select the larger of those specimens (NMW 46682:1, 93 mm SL) as the lectotype of Eu- anemus longipinnis. Four specimens deposited in MCZ (orig- inally MCZ 9834 (4 specimens), now di- vided between MCZ 9834 (3 specimens) and MCZ 33450 (1 specimen)) were iden- tified as syntypes of Euanemus longipinnis by Eigenmann & Eigenmann (1890:292) in their redescription of Epapterus dispilurus. However, given that all specimens cited as syntypes in the original description of Eu- anemus longipinnis are accounted for in the NMW holdings, we do not consider the MCZ material to be part of the type series. Status of Epapterus chaquensis In the abstract of their original descrip- tion of Epapterus chaquensis, Risso & Ris- so (1962:4) noted that their nominal species differed from E. dispilurus in the form of the dark humeral spot and in the presence of dark pigmentation of the ventral (=pel- vic) fin. Within the main body of the text Risso & Risso (1962:6) cited, however, a more extensive series of characters discrim- inating the two species, proposing that E. chaquensis differed from E. dispilurus “principally in the dark coloration of the anal fin and basal and distal regions of the ventrals [=pelvic fins]. The humeral mark is not rounded but obliquely lengthened. Lesser number of rays in the anal and pec- toral. Head and eyes smaller, etc.’ (our translation). Risso & Risso gave no indication that PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON they examined comparative specimens of E. dispilurus, rather their literature cited and introductory remarks indicate that their comparisons were based on literature infor- mation, in particular the original description of E. dispilurus (Cope, 1878:677) and the line drawing of the lectotype of that species included in Fowler’s listing of the fishes of Peru (1941, 1945). Cope (1878), in his orig- inal description of E. dispilurus (1878), commented neither on the form of the hu- meral spot nor made mention of any pig- mentation of the pectoral, pelvic, and anal fins. Fowler’s relatively simple line drawing (1941:468, fig. 26; 1945:66, fig. 26) was based on the lectotype of E. dispilurus, a specimen which is now in very poor con- dition and which may not have been in a much better state at the time that Fowler prepared the figure. Specimens of other spe- cies cited by Cope (1878) were already in poor condition less than two decades later when reported on in 1906 by Fowler (Vari 1992:117-—118). More recently collected specimens of E. dispilurus from the Peru- vian Amazon, the type region of that spe- cies, have a humeral spot which is much more horizontally elongate than that shown in Fowler’s figure of the species. The hu- meral pigmentation pattern in E. dispilurus specimens from the Peruvian Amazon is comparable to that in population samples of Epapterus from the Rio Paraguay, the type- region for E. chaquensis. Similarly, Fowl- er’s drawing (1941, 1945) does not include the dark anal-, pelvic-, and pectoral-fin pig- mentation which is typical for Amazonian Epapterus populations samples and which also occurs in population samples of the ge- nus from the Rio Paraguay basin as noted by Risso & Risso. The incomplete repre- sentation of the dark body and fin pigmen- tation in Fowler’s figures (1941, 1945) pre- sumably lead Risso & Risso to erroneously propose that the Amazonian and Paraguay- an populations differed in details of dark pigmentation. Our comparisons of popula- tion samples from the two basins have, in contrast, failed to reveal any consistent dif- VOLUME 111, NUMBER 4 Table 1.—Ranges in population samples of Epap- terus from the Rio Paraguay and Rio Amazonas basins for meristic and morphometric features cited by Risso & Risso (1962) as distinguishing EF. dispilurus (Rio Amazonas) from E. chaquensis (Rio Paraguay). Basin Feature Rio Amazonas Rio Paraguay Anal-fin rays 52-62 53-61 Pectoral fin rays 11-13 12-13 Head length in SL 18.3—21.9 18.5—20.9 Eye diameter in HL 25.0—-33.1 26.7—29.8 ference in these or any other pigmentation patterns. Risso & Risso (1962:6) also cited pur- ported differences in the numbers of anal- and pectoral-fin rays and relative sizes of the head and eye as distinguishing Epap- terus chaquensis from E. dispilurus. Pre- sumably their comparative data for the lat- ter species was taken from the original de- scription by Cope (1878) which was based on only two specimens. Similarly, Risso & Risso had only five specimens of E. cha- quensis available for their analysis. A com- parison of larger population samples from the Amazon and Paraguay basins (see “‘Ma- terial Examined’) has shown that the de- gree of variation in the number of anal- and pectoral-fin rays and in the range in the rel- ative size of the head and eyes in the pop- ulation samples from the Amazon and Par- aguay basis is significantly greater than in- dicated by the limited type-series of E. dis- pilurus and E. chaquensis. As a consequence there is broad, or complete, overlap be- tween the samples from the two basins in all features proposed by Risso & Risso (1962:6) to distinguish the nominal species (Table 1). We have been unable to identify any meristic, morphometric, or color pat- tern differences that would justify the con- tinued recognition of E. chaquensis and we consequently place that species into the synonymy of E. dispilurus. Material examined.—Total specimens: 75. Specimens from which counts and mea- surements were taken: 50, 59-123 mm SL. 1003 Argentina: Chaco: Resistencia, Laguna along side of the Rio Salado (27°27'S, 58°59'W), MLP 8017, 1 (124). Rio Negro, Resistencia (27°27'S, 58°59'W), MZUSP 10252—53, 2 (88—93). Bolivia: El Beni: Canal San Gregorio, Trinidad (14°47'S, 64°47'W), MZUSP 27816, 2 (63-72). Trinidad (14°47’S, 64°47'W), USNM 278586, 3 (59-92). Rio Itenez, Londra, USNM 278563, 2 (78-100). Rio Matos, 48 km E San Borja, below road crossing (14°55’S, 66°17’W), USNM 305651, 2 (79-102). Brazil: Amazonas: Rio Javari of Rio So- lim6es (4°21'S, 70°02'W), MCZ 9834, 3 (75—107; erroneously identified as syntypes of Euanemus longipinnis by Eigenmann & Eigenmann 1888:292); MCZ 33450, 1 (78; erroneously identified as syntype of Euane- mus longipinnis by Eigenmann & Eigen- mann 1888:292). Humaita, Igarapé Joari (7°31'S, 63°02’W), MZUSP 27912, 2 (88— 95). Rio Aripuana, Igarapé on Ilha do Cas- tanhal (~5°07'S, 60°24’W), MZUSP 48866, 1 (73). Hyavary [=Rio Javari] (4°21'S, 70°02’W), NMW 46682:1, 1 (93, lectotype of Euanemus longipinnis); NMW 46682:2 (79, paralectotype of Euanemus longipinnis). Lago Terra Preta, Januari (3°12'S, 60°05’W), USNM 261422, 2 (73- 86). Mato Grosso: Lagoa on Fazenda Ong¢a Magra, Municipio de Caceres (~16°04’S, 57°41'W), MZUSP 38172, 1 (90). Mato Grosso do Sul: Rio Aquidauana, Baia da Oncga or Jatobd, Fazenda Alegrete (~19°44'S, 56°50'W), MZUSP 40084, 4 (98-122). Paraguay: Cordillera: Rio Piribebuy, 5 km N of Emboscada (~25°29'S, 57°03'W), MHNG 2212.27, 1 (123). Presidente Hayes: Rio Negro, 6 km S of Chaco-I (~25°15'S, 57°38'W), MHNG 2212.30, 1 (116). Rio Paraguay at San Antonio, MHNG 2213.25, 1 (119). Rio Aguaray-guazu, at km 117 of Transchaco Road (24°47’S, 57°19'’W), MHNG 2213.26, 1 (123). Rio Negro at Trans-Chaco Highway crossing (~25°10'S, 58°30'W), USNM 232304, 1 (87). Peru: Peruvian Amazon, ANSP 21353, 1 1004 (107; lectotype of Epapterus dispilurus), ANSP 21354, 1 (96, paralectotype of Epap- terus dispilurus). Loreto: Rio Ucayali basin, Contamana (7°20'S, 75°01’W), ANSP 103412, 1. Reserva Natural Pacaya-Sami- ria, Cano Ungurahue of Rio Pacaya (4913'S, 74°24'W), MUSM 2524, 1 (66). Ucayali: Rio Ucayali, Provincia Coronel Portillo, Bagazan, MZUSP 26183, 4 (96— 110). Provincia Coronel Portillo, Pucallpa, Cashibococha (7°33'S, 74°53'W), MZUSP 26318, 6 (100-109). Provincia Coronel Portillo, Pucallpa, Rio Ucayali (8°23’S, 74°32'W), MZUSP 26410, 7 (51-63). Prov- incia Coronel Portillo, Rio Aguaytia, Nueva Requena (8°20’S, 74°34’W), USNM 261388, 6 (59-106). Provincia Coronel Portillo, Rio Ucayali at Pucallpa (8°23’S, 74°32'W), USNM 263114, 1 (59). Provin- cia Coronel Portillo, Rio Ucayali, Utuquinia (8°15’S, 74°33'W), USNM 263115, 4 (83- 89); USNM 273591, 5 (103-115); USNM 273615, 4 (115-128). Epapterus blohmi Vari et al., 1984 Epapterus blohmi Vari et al., 1984:463, fig. 1 (type locality: Venezuela, Guarico State, Fundo Pecuario Masagural, Laguna Los Gudcimos).—Machado-Allison & Moreno, 1993:87 (Venezuela, Guarico State, Rio Orituco).—Machado-Allison et al., 1993:130 (Venezuela, lower Ilan- os).—Lasso et al., 1995:4, 7, 8 (Venezue- la, flooded Nanos; ecology).—Taphorn et al., 1997:83 (Venezuela). Publications dealing with aspects of the distribution and ecology of Epapterus bloh- mi which appeared subsequent to its origi- nal description are cited in the synonymy for the species. Remarks.—Vari et al. (1984) provide a detailed description of Epapterus blohmi and there is no need to repeat that account in its entirety. Nonetheless, in the course of this study we had the opportunity to ex- amine a greater number of specimens of E. blohmi from a wider geographic range (see “Distribution”? below) than were available PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to Vari et al. (1984). This increased sample, not unexpectedly, revealed that the follow- ing meristic values were found to demon- strate a broader range in this study than re- ported in the original description of E. bloh- mi: dorsal-fin rays 2 or 3; pectoral-fin rays 9 to 11; pelvic-fin rays 13 to 16; anal-fin rays 49 to 61; gill-rakers 29 to 40; precau- dal vertebrae 14 or 15; caudal vertebrae 34 to 36; total vertebrae 49 to 51 (note: ver- tebral counts presented in original account did not include consolidated elements in Weberian apparatus). Vari et al. (1984) re- ported that teeth were absent on the lower pharyngeal tooth-plate of FE. blohmi. Ex- amination of additional cleared and stained specimens has revealed that the cited ab- sence was an individual variant in a smaller specimen and that dentition on the lower pharyngeal tooth-plate is typically present in the species. Distribution.—Vari et al. (1984) de- scribed Epapterus blohmi on the basis of a series of specimens collected from Guarico State, Venezuela, in the central portions of the Rio Orinoco basin. Subsequent collect- ing efforts elsewhere in that river system have shown that the species is widespread in Apure State and occurs in the Portuguesa State, both of which are located to the west of the type-locality. These efforts have also revealed that the range of the species ex- tends east into the Anzoategui State. More unexpected was the discovery of EF. blohmi in the Rio Tuy system, a Caribbean versant basin along the northern coast of Venezue- la, a distinct range extension to the north of the type locality of the species and the first reported occurrence of E. blohmi outside of the Rio Orinoco basin. A single lot of E. blohmi examined during this study origi- nated in Quebrada Caraballo within the Rio Tuy system, Miranda state. The fishes of the Rio Tuy basin are still poorly known, a problem exacerbated by the large scale pollution of the main chan- nel of the river. Nonetheless, a pattern has become apparent in recently revised groups in which species are found to be common VOLUME 111, NUMBER 4 to the Rio Tuy and Rio Orinoco basins. In addition to Epapterus blohmi, these include the characiform species Creagrutus melas- ma (Vari et al. 1994:95) and Steindachner- ina argentea (Vari 1991:41). Although the phylogenetic relationships of most species endemic to the Rio Tuy basin remain un- certain, there is some indication of sister- group relationships between species in that basin and those of the Rio Orinoco system (e.g., Serrasalmus neveriensis Machado-Al- lison et al., 1993:53; Machado-Allison & Fink, 1996:113). The distribution data from Epapterus blohmi is congruent with the hy- pothesis of a once continuous ichthyofauna between the Rio Tuy and Rio Orinoco ba- sins which was disrupted by the uprise of the Serrania de la Costa as proposed by Ma- chado-Allison et al. (1993:53). Material examined (in addition to that cited in Vari et al. 1984).—Venezuela: An- zoategui. Rio Zuata, near Zuata (8°22'N, 65°22'W), USNM 316822, 4. Apure. Mod- ulo UNELLEZ, near Cafio Caicara, MCNG 3599, 8. Modulo UNELLEZ, east dyke, MCNG 2435 1; MCNG 3932, 1. Modulo de Mantecal, MCNG 19593, 1. Hato El Frio (7°49'N, 68°54'W), MCNG 5963, 1; MCNG 9914, 1; MCNG 9668, 10. Laguna El Pozon, MCNG 24075, 1. Miranda. Que- brada Caraballo, along highway between Caracas and Caucagua, parcela Yaguara (Rio Tuy basin), MCNG 27358, 7 (of 44). Portuguesa. Cafio Maraca (8°47’'N, 69°05’W), MCNG 8637, 1. Districto Turen, Cano Sal- ado ~5 km S of Nueva Florida (latter lo- cality at 8°57'N, 69°01’W), MCNG 12679, 1. Acknowledgments Research at museums in North and South America and field work associated with this project was supported, in part, by the Neo- tropical Lowland Research Program of the Smithsonian Institution. We thank the fol- lowing individuals and institutions for loans of specimens, hospitality during visits, and other types of assistance: Scott A. Schaefer and William Saul (ANSP); Donald Taphorn 1005 (MCNG); Sonia Muller (MHNG); Adriana Almir6n and Mercedes Azpelicueta (MLP); Hernan Ortega and Fonchii Chang (MUSM); José Lima de Figueiredo, Heral- do Britski, Naércio A. Menezes, Osvaldo T. Oyakawa, and Mario de Pinna (MZUSP); and Ernst Mikschi and Barbara Herzig (NMW). Technical assistance at USNM was ably provided by Sandra Raredon and Jeffrey Clayton. Figures 1 and 2 were pre- pared by T. Britt Griswold. The Portuguese translation of the abstract was generously provided by Mario C. C. de Pinna. The pa- per benefitted from the comments and sug- gestions of Stanley H. Weitzman, Jonathan W. Armbruster, Michael E. Retzer, and Scott A. Schaefer. Literature Cited Bohlke, E. B. 1984. Catalog of type specimens in the ichthyological collection of the Academy of Natural Sciences of Philadelphia——The Acad- emy of Natural Sciences of Philadelphia, Spe- cial Publication 14:i—viii, 1-246. Bohlke, J. 1951. Description of a new auchenipterid catfish of the genus Pseudepapterus from the Amazon basin.—Stanford Ichthyological Bul- letin 4(1):38—40. Burgess, W. E. 1989. An atlas of freshwater and ma- rine catfishes. A preliminary survey of the Sil- uriformes. T. EF H. Publications, Inc. Neptune City, New Jersey, 784 pp. Cope E. D. 1878. Synopsis of the fishes of the Peru- vian Amazon obtained by Professor Orton dur- ing his expeditions of 1873 and 1877.—Pro- ceedings of the American Philosophical Society 17(101):673—701. Cuvier, G., & A. Valenciennes. 1840. Histoire naturelle des poissons. Ch. Pitois, Paris, i-xviii, 1—504. Eigenmann, C. H. 1910. Catalogue of the fresh-water fishes of tropical and south temperate Ameri- ca.—Reports of the Princeton University Ex- peditions to Patagonia, 1896—1899 3:375—-511. , & W.R. Allen. 1942. Fishes of western South America. Lexington, The University of Ken- tucky, i-xv, 1—494. , & R. S. Eigenmann. 1888. Preliminary notes on South American Nematognathi, I—Proceed- ings of the California Academy of Sciences, 2nd. Series 1(2):119—172. , . 1890. A revision of the South American Nematognathi or cat-fishes.—Occa- sional Papers of the California Academy of Sci- ences 1:1—508. 1006 Eschmeyer, W. N. 1990. Catalog of the genera of Re- cent fishes. California Academy of Sciences, San Francisco l—v, 1—697. Ferraris, C. J., Jr. 1988. The Auchenipteridae: putative monophyly and systematics with a classification of the neotropical doradoid catfishes (Ostario- physi: Siluriformes). Unpublished Ph.D. disser- tation, City University of New York. , & R. P Vari. 1999. The South American cat- fish genus Auchenipterus (Ostariophysi: Siluri- formes: Auchenipteridae): monophyly and re- lationships, with a revisionary study.—Zoolog- ical Journal of the Linnean Society (in press). Fowler, H. W. 1906. Further knowledge of some Het- erognathus fishes, Part. Proceedings of the Academy of Natural Sciences of Philadelphia 58:293-351. . 1914. Fishes of the Rupununi River, British Guiana.—Proceedings of the Academy of Nat- ural Sciences of Philadelphia 66:229—284. . 1915. Notes on Nematognathous fishes.—Pro- ceedings of the Academy of Natural Sciences of Philadelphia 67:203—243. . 1940. A collection of fishes obtained by Mr. William C. Morrow in the Ucayali River basin, Peru.—Proceedings of the Academy of Natural Sciences of Philadelphia 91(for 1939):219—289. . 1941. Los peces del Pert. Catalogo sistema- tico de los peces que habitan en aquas peruanas (Continuacion).—Boletin del Museo de Historia Natural “‘Javier Prado” Lima 5(19):466—487. . 1945. 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Serrasalmus neveriensis una nueva especies de caribe de Venezuela y redescripcio de Serra- salmus medinai Ramirez, 1965.—Acta Biolo- gica Venezuelica 14(4):45—60. , & H. Moreno. 1993. Estudios sobre la com- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON munidad de peces del Rio Orituco, Estado Guarico, Venezuela. Parte I. Inventario, abun- dancia relativa y diversidad.—Acta Biologica Venezuelica 14(4):77—94. , E Mago-Leccia, O. Castillo, R. Royero, C. Marrero, C. Lasso, & E Provenzano. 1993. Lis- ta de especies reportadas en los diferentes cuer- pos de agua de los bajos llanos de Venezuela. Anexo I. Pp. 129-136 in A. Machado-Allison, Los peces de los llanos de Venezuela: un ensayo sobre su historia natural. Second Edition. Univ- ersidad Central de Venezuela, Caracas. Miranda-Ribeiro, A de. 1918. Tres generos e dezesete especies novas de peixes Brasileiros, determi- nados nas collecgoes do Museu Paulista.—Re- vista do Museu Paulista 10:629—646. Mees, G. FE 1974. 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Part 1: the drift- wood catfishes.—Tropical Fish Hobbyist 34(10):26—45. Steindachner, FE 1881. Beitraége zur Kenntnis der Fluss- fische Stidamerika’s IIJ].—Denkschriften der Akademie der Wissenschaften, Mathematisch- Naturwissenschaftliche Klasse, Wien 44[for 1882]:1-17. . 1882. Beitrage zur Kenntnis der Flussfische Siidamerika’s [1V.—Denkschriften der Akademie der Wissenschaften, Mathematisch-Naturwis- senschaftliche Klasse, Wien 46:1—44. . 1915. Vorlaufiger Bericht tiber einige neue Stisswasserfische aus Siidamerika.—Anzeiger der Akademie der Wissenschaften, Wien 52: 199-202. Taphorn, D. C., R. Royero, A. Machado-Allison, & F Mago-Leccia. 1997. Lista actualizada de los pe- ces de agua dulce de Venezuela. Pp. 55—100 in E. La Marca, ed., Vertebratos actuales y fosiles VOLUME 111, NUMBER 4 1007 de Venezuela. Serie Catalogo Zooldgico de tariophysi).—Smithsonian Contributions to Zo- Venezuela. Volume 1. Museo de Ciencia y Tec- ology 529:i-iv, 1-137. nologia de Mérida, Merida, Venezuela. , A. S. Harold, & D. C. Taphorn. 1994. Crea- Troschel, E H. 1882. Bericht tiber die Leistungen in grutus melasma, a new species of characid fish der Ichthyologie wahrend des Jahres 1881.— (Teleostei: Characiformes) from upland streams Archiv fiir Naturgeschichte 48(5):431—456. of northern Venezuela.—Proceedings of the Bi- Vari, R. P. 1991. Systematics of the neotropical char- ological Society of Washington 107(1):90—96. aciform genus Steindachnerina Fowler (Pisces: , S. L. Jewett, D. C. Taphorn, & C. R. Gilbert. Ostariophysi).—Smithsonian Contributions to 1984. A new catfish of the genus Epapterus Zoology 507:i-iv, 1-118. (Siluriformes: Auchenipteridae) from the Ori- . 1992. Systematics of the neotropical characi- noco River basin.—Proceedings of the Biolog- form genus Cyphocharax Fowler (Pisces: Os- ical Society of Washington 97(2):462—472. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 111(4):1008—1015. 1998. Two new subspecies of Cinnycerthia fulva (Aves: Troglodytidae) from the southern Andes J. V. Remsen, Jr. and Robb T. Brumfield Museum of Natural Science, Louisiana State University, Baton Rouge, Louisiana 70803, U.S.A.; (RTB) Present address: Laboratory of Molecular Systematics, National Museum of Natural History, Smithsonian Institution MRC 534, Washington, D.C. 20560-0534, U.S.A. Abstract.—Analysis of geographic variation of plumage pattern and color within Cinnycerthia fulva (Troglodytidae) of the southern Andes revealed that this taxon, formerly treated as monotypic, consists of three discrete units. We describe two new subspecies; one (C. f. fitzpatricki) from the isolated Cordillera Vilcabamba, depto. Cuzco, Peru, and the other (C. f gravesi) from depto Puno, Peru, south to depto. Cochabamba, Bolivia. Our previous analysis of geographic var- iation in size and plumage color of Cinny- certhia wren populations of the Andes (Brumfield & Remsen 1996) suggested that the form previously treated as a single spe- cies, Cinnycerthia peruana (Sepia-brown Wren; Troglodytidae) consists of at least three biological species: C. olivascens of the northern Andes south to northern Peru, C. peruana of the Andes of Peru from dep- to. Amazonas south to the Rio Apurimac canyon, and C. fulva, from depto. Cuzco, Peru, to the Andes of Bolivia. We also not- ed that C. fulva contains at least three dis- tinctive populations, only one of which is named: a dark-crowned population restrict- ed to the isolated Cordillera Vilcabamba, depto. Cuzco, Peru; a population in the main Andes of southern depto. Cuzco, in- cluding the type locality of fulva; and a small-sized, pale population in the main Andes from southern depto. Puno to depto. Cochabamba, Bolivia. We herein formally name the two undescribed populations. The Cordillera Vilcabamba, depto. Cuz- co, Peru, is a mountain range isolated from the main chain of the Andes by deep river valleys with tropical, non-montane habitats (Terborgh 1971, Haffer 1974). Although specimens of birds collected there by John Weske and John Terborgh have yet to be analyzed in a systematic manner, some en- demic taxa have been described or are in the process of being described: Schizoeaca vilcabambae Vaurie et al., 1972 (see Rem- sen 1981), Cranioleuca marcapatae weskei Remsen, 1984, Atlapetes rufinucha terbor- ghi Remsen, 1993, and undescribed subspe- cies of Ochthoeca fumicolor and Coeligena violifer. Our analysis of geographic variation in the Cinnycerthia peruana superspecies (Brumfield & Remsen 1996) revealed that the population from the Cordillera Vilca- bamba has a highly distinctive head pattern (Fig. 1) approached by no other population in the complex. This population, which is part of the complex that Brumfield & Rem- sen (1996) treated as a separate species, C. fulva, may be known as: Cinnycerthia fulva fitzpatricki, new subspecies Holotype.—American Museum of Natu- ral History (AMNH) 820311; mist-netted by John S. Weske and John W. Terborgh on 11 Jul 1967 at Cordillera Vilcabamba, elev. 2090 m, depto: Cuzco, Peru, 12738553 VOLUME 111, NUMBER 4 1009 Fig. 1. gravesi. 73°36'W. The specimen, prepared by Weske (#1285), is a male (largest testis 2 % mm diam) with a completely pneumatized skull and little fat. Paratypes.—Six other specimens were also mist-netted in the Cordillera Vilcabam- ba by Weske and Terborgh and prepared by Weske: AMNH 820059, 14 km E. of Lou- Head patterns, from top to bottom, of Cinnycerthia peruana, C. f. fitzpatricki, C. f. fulva. and C. f- isiana, 2050 m, 12°39'S, 73°34'W, 6, 12 Aug 1966; AMNH 820210, 1740 m, 12°38'S, 73°38’W, ¢ (skull “‘scarcely pneu- matized’’), 12 Aug 1967; AMNH 820211, 1750 m, 12°38'S, 73°36'W, ¢ (skull com- pletely pneumatized), 28 Jun 1968; AMNH 820309, 2160 m, 12°38'S, 73°36'’W, (skull pneumatization incomplete), 29 Jun 1010 1967; AMNH 820310, 2200 m, 12°38’S, 73°36'W, 3 (skull pneumatization com- plete), 30 Jun 1967; and (6) AMNH 820507; 23830r mas 2 Si SSS Wane Jul 1967. Diagnosis.—Cinnycerthia f. fitzpatricki can be distinguished at a glance from any other taxon in the genus by the narrow blackish border of the forecrown and the dark crown that contrasts with the back (Fig. 1). In Cinnycerthia peruana, the fore- crown is slightly paler, not darker, than the rest of the crown and the crown and back are the same color. In C. f. fulva, the crown and back are concolorous. Otherwise, fi- tzpatricki is similar to C. f. fulva but has a whitish, not light brown, superciliary, and it is paler below (but sample sizes of both perhaps too small for confidence on this character). As noted by Brumfield & Rem- sen (1996), this subspecies may be larger in linear measurements than the other subspe- cies of C. fulva, but sample sizes are too small for statistical testing. Description.—Capitalized color names are from Ridgway (1912). Forecrown nar- rowly Blackish Brown, blending to dark brown (near Raw Umber) crown, which then blends to a uniformly rich brown nape and back (Proutt’s Brown). Uppertail co- verts slightly more reddish than back. Outer webs of remiges (and both webs of inner secondaries) rich brown strikingly banded with narrow blackish bars; inner webs of most remiges dark fuscous. Upper wing co- verts rich brown, faintly banded blackish; underwing coverts rich brown. Rectrices rich rufous brown strikingly banded with narrow blackish bars, slightly less conspic- uously than remiges. Conspicuous buffy whitish superciliary from nares over eye to neck, sharply demarcated from the crown, which tends to be darkest at border of su- perciliary, and from broad, dark brown (Raw Umber) postocular stripe. Narrow whitish eyering. Lores brown, not distinctly separated from superciliary. Lower portion of auricular below postocular area dull light brownish (near Cinnamon), distinctly de- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON marcated from postocular area. Malar re- gion same color as lower auriculars. Chin and throat buffy whitish, noticeably paler than rest of underparts and malar area but not distinctly demarcated from them. Breast brown (Sayal Brown), blending to more ru- fous brown belly. Lower flanks, thighs, and undertail coverts darker (Proutt’s Brown) than rest of underparts, as dark as back. Soft part colors recorded by Weske: iris brown; bill mainly black, shading to pale gray on gonys; legs brownish-gray. Mea- surements (mm): wing chord = 59.5; ex- posed culmen = 12.3; tail length = 55.8; tarsus length = 21.2. Variation.—The seven specimens from the Vilcabamba show little individual vari- ation except that one (AMNH 820211) has a slightly darker buffy chin, throat, and su- perciliary. One young bird (skull scarcely pneumatized; AMNH 820210) matches adult individuals closely in plumage, but its mandible is noticeably paler, a pale yellow- ish horn in the dried specimen (and de- scribed as “yellow-brown” by its prepara- tor); those of the other birds are blackish. Measurements also show little variation (Table 1). Natural history.—The only information available comes from the specimen labels. All specimens were mist-netted in humid, montane cloudforest and elfin forest, from 1740 to 2830 m, a considerable elevational range. Weske (1972) gave its elevational limits as 1710 to 2830 m. A male collected on 29 Jun had enlarged testes (left 6 mm, right 4.5 mm). A female collected on 12 Aug may have been in breeding condition (ovary 7 mm, largest ovum 1.2 mm). The other specimens did not have enlarged go- nads. Etymology.—It is a pleasure to name this distinctive taxon for John W. Fitzpatrick, whose fieldwork in depto. Cuzco and else- where in southern Peru has contributed greatly to knowledge of South American birds. His recent specimens from near the type locality of C. fulva (see below) were VOLUME 111, NUMBER 4 1011 Table 1—Measurements (in mm) of Cinnycerthia fulva fitzpatricki, C. f. fulva, and C. f. gravesi southern Peru and northern Bolivia. The numbers in the columns are the means followed by the ranges in parentheses. Population (n) C. f. fitzpatricki (2) C. f. fulva (2) C. f. gravesi (depto C. f; gravesi (depto C. f. gravesi (depto C. f. fitzpatricki (4) C. f. fulva (0) C. f. gravesi (depto C. f. gravesi (depto . Puno; 2) . La Paz; 5) . Cochabamba; 3) . Puno; 5) . La Paz; 7) Wing chord Exposed culmen Males 60.8 (59.5—62.0) 12.6 (12.3—13.2) 57.2 (56.7—58.6) 13.5 (13.4—13.5) 59.3 (57.8-60.7) 12.8 (12.4-13.1) 57.2 (56.1-58.3) 12.9 (12.0-13.8) 56.7 (54.8-58.8) 12.1 (11.2—13.6) Females 55.7 (52.8—-57.2) 12.8 (12.4—12.8) 56.3 (54.1-58.8) 12.4 (11.2—13.1) 55.0 (54.2—56.0) 12.9 (12.0—-13.8) C. f. gravesi (depto. Cochabamba; 2) 53.3 (52.1—54.4) 12.0 (12.0—12.0) Tail length 55.6 (55.1—55.8) 55.6 (54.2—57.0) 54.7 (53.5—55.8) 54.6 (53.1—56.5) 54.9 (53.0—56.7) 54.1 (51.8-56.9) 52.5 (50.3—55.4) 54.6 (53.1—56.5) 53.5 (53.1—53.9) Tarsus length 21.5 (21.1—21.6) 22.8 (21.1—24.4) 20.7 (20.4—21.0) 22.0 (21.1—23.4) 21.0 (20.9-21.0) 20.7 (19.7—21.4) 20.8 (19.9-21.7) 22.0 (21.1—23.4) 20.4 (19.8—21.0) critical to the taxonomic conclusions of this paper. Discussion.—Does the distinctive crown of fitzpatricki merit recognition as a spe- cies? Because its dark crown is a unique character within the genus Cinnycerthia, some might treat this taxon as a species. Because fitzpatricki is completely isolated from other populations of Cinnycerthia, di- rect assessment of reproductive isolation from other populations is not possible. However, we cannot find any comparable situation in the Troglodytidae in which two species-level taxa (e.g., in Hellmayr 1934, Ridgely & Tudor 1989, Fjeldsa & Krabbe 1990) differ only in crown color and are otherwise extremely similar in plumage. Therefore, we recommend that fitzpatricki be treated as a subspecies of the biological species C. fulva in the absence of infor- mation on potential reproductive isolating mechanisms, particularly vocalizations. Al- though geographically closer to some pop- ulations of C. peruana, we place it within C. fulva because of the plumage similari- ties, particularly the prominent superciliary; also, most specimens of fitzpatricki clearly fit within C. fulva on the basis of measure- ments (Brumfield & Remsen 1996). We previously proposed (Brumfield & Remsen 1996) that the remaining popula- tions of C. fulva can be divided into two discrete units, the population in depto. Cuz- co (where the type locality is) and the pop- ulation from depto. Puno, Peru, south through depto. La Paz, Bolivia, to depte. Cochabamba. As discussed by Brumfield & Remsen (1996), few specimens exist from depto. Cuzco. The two that we examined, collected by J. Weske and J. Fitzpatrick at Pillahuata, were collected approximately 15 km north of the type locality of C. fulva. They match Sclater’s type description (“‘brown, pale, but ill-defined superciliary mark’’) in the key feature that we propose distinguishes the Cuzco population from those to the south, namely that the super- ciliary is not dull white, but ochraceous brown. We have yet to find a specimen from farther south with a superciliary that is not at least partially whitish. Also, the auriculars of the specimens from Pillahuata are darker than those from farther south, thereby reducing the contrast between it and the dark brown postocular stripe. Hellmayr (1934), who examined the unique type of fu/va in the British Museum, stated that it “is more reddish, less oliva- ceous, throughout than a Bolivian series. The divergency requires confirmation by additional specimens.’’ We are unable to see any such differences between the Pil- lahuata specimens and specimens from far- ther south. Curiously, Hellmayr did not re- 1012 mark upon the light brown superciliary of the type specimen, although he emphasized the “‘well-defined, large, buffy white super- ciliary streak’’ as a character for distin- guishing C. fulva from C. peruana, which applies primarily to specimens that he also examined from Bolivia. The Weske-Fitzpatrick specimens are the only two from depto. Cuzco, other than the type specimen, that we have been able to locate. Although a larger sample would be desirable, we believe that the distinctive features of these specimens signal that the Cuzco population is a separate taxonomic unit, and that they do not represent individ- ual or clinal variation, especially given the relative uniformity (other than individual variation in number of white facial feathers) of the southern population over a range of at least 550 km. Therefore, we propose to name the southern population: Cinnycerthia fulva gravesi, new subspecies Holotype.—Academy of Natural Scienc- es of Philadelphia (ANSP) 138618; ¢ col- lected at Incachaca, 8000 ft, depto. Cocha- bamba, Bolivia, on 4 Jun 1937 by M. A. Carriker, Jr. Paratypes.—Peru (depto, Puno): Santo Domingo, 6000 ft (AMNH 146335; 6, 14 Sep 1916; H. Watkins); Santo Domingo, 4500 ft (AMNH 502123; ¢, 24 May 1901; G. Ockenden); Limbani, 9500 ft (AMNH 502122; 2, 28 Feb 1904; G. Ockenden); Valcé6n, 5 km NNW Quiaca, 3000 m (Lou- isiana State University Museum of Natural Science [LSUMZ] 98604-607; 11-24 Oct 1980; L. C. Binford, M. Sanchez S., T. S. Schulenberg); Abra de Maruncunca, 10 km SW San Juan del Oro, 2000 m (LSUMZ 98608; 26 Nov 1980; T. S. Schulenberg). Bolivia (depto, La Paz): Rio Zongo Valley, 1.9 road km downstream Saenani, 2236 m (Delaware Museum of Natural History 67191; 36, 15 Mar 1979; D. C. [Cole] Schmitt); Sandillani, 6800 ft (ANSP 121196-197; males, 13 and 19 Dec 1934; PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON M. A. Carriker, Jr.); Sacramento Alto, 2575 m (LSUMZ 90885-890; 27-31 Jul 1979; L. S. Hale and J. V. Remsen); ca. 1 km S Chuspipata, 2725 and 3050 m (LSUMZ 102809-816; 30 Jul-5 Aug 1981; S. M. Lanyon, J. V. Remsen, T. S. Schulenberg, D. A. Wiedenfeld). Bolivia (depto, Cocha- bamba, prov. Chapare): Incachaca, 11000 ft (ANSP 138617; 3d, 10 Jun 1937; M. A. Carriker, Jr.); 10000 ft (ANSP 138616; 2, 1 Jun 1937; M. A. Carriker, Jr.); 2600 m (Carnegie Museum of Natural History [CM] 85205, 85234; 25 Mar and 5 Apr 1921; J. Steinbach); 2500 m (CM 85176; 28 Feb- 1921:-J. Stembach)e2200 tin (LSUMZ 38048; 17 (?) Nov 1921; J. Stein- bach); 2225 m (CM 120415; 7 Sep 1927; J. Steinbach); El Limbo, 2200 m (LSUMZ 36405; 14 Oct 1960; E Steinbach). Diagnosis.—Cinnycerthia f. gravesi can be distinguished from C. f. fulva by its whitish, rather than buffy brown (close to Clay Color), superciliary stripe and fore- crown (Fig. 1). It also is paler below than C. f: fulva (but only two of the latter are available for comparison). In having a whit- ish superciliary that contrasts strongly with darker auriculars, the head pattern resem- bles that of distant C. f. fitzpatricki more so than adjacent C. f. fulva. Description.—Forecrown mostly dull white, mixed with two or three brown feathers. Crown and rest of dorsum rich brown (near Proutt’s Brown). Uppertail co- verts slightly more reddish than back. Outer webs of remiges (and both webs of inner secondaries) rich brown strikingly banded with narrow blackish bars; inner webs of most remiges dark fuscous. Upper wing co- verts rich brown, faintly banded blackish; under-wing coverts rich brown. Rectrices rich rufous brown strikingly banded with narrow blackish bars, slightly less conspic- uously than remiges. Conspicuous buffy whitish superciliary extends from nares over eye to neck and becomes darker pos- teriorly; it is sharply demarcated from the crown, which tends to be slightly darker at border of superciliary, and from broad, rich VOLUME 111, NUMBER 4 brown (near Proutt’s Brown) postocular stripe. Narrow whitish eyering. Lores brown mixed with white. Lower portion of auricular below postocular area is dull light brownish (near Cinnamon), distinctly de- marcated from postocular area. Malar re- gion same color as lower auriculars. Chin and throat buffy whitish, noticeably paler than rest of underparts and malar area but not distinctly demarcated from them. Breast brown (Sayal Brown), blending to more ru- fous brown belly. Lower flanks, thighs, and undertail coverts darker (Proutt’s Brown) than rest of underparts, as dark as back. Soft part colors recorded by Carriker: iris gray; bill black, bluish black below; legs dark horn. Measurements (mm): wing chord = 57.0; exposed culmen = 11.6; tail length = 56.7; tarsus length = 20.9. Variation.—As noted by Brumfield & Remsen (1996), about 30% of the individ- uals examined show “extra” white in the face, similar to but not nearly as extensive as variation in C. peruana (Graves 1980). Otherwise, the series is relatively uniform in plumage color and pattern, with individ- uals from depto. Puno virtually identical to individuals from depto. Cochabamba. Also, specimens taken in 1980 from depto. Puno are virtually identical in plumage to those taken there from 1904 to 1916, and so we see no evidence for “‘foxing.’’ However, the recent specimens from July and August from Chuspipata and Sacramento Alto, dep- to. La Paz, are puzzling. These stand out strongly as being less reddish above and be- low, paler ventrally, and darker dorsally. Although we previously attributed this to seasonal wear, we now think that the dif- ferences are too great to be caused by wear, and we found no seasonal effects on plum- age color in deptos. Puno or Cochabamba. Furthermore, the plumage of these puzzling specimens does not seem to be more abrad- ed than specimens taken elsewhere at other times of the year. We would be tempted to describe the depto. La Paz population as a distinct subspecies, but two specimens from Sandillani, fewer than 10 km away and in 1013 the same drainage, are virtually identical to specimens from elsewhere in the range. The basis for this variation can only be resolved by further field-work in the area. Natural history.—The scant information published concerning the natural history of Cinnycerthia peruana (e.g., Ridgely & Tu- dor 1989, Fjeldsa & Krabbe 1990) probably applies in general to C. f gravesi. The only published information that applies explicit- ly to C. f. gravesi comes from Remsen (1985), who classified it as a rare insecti- vore at a study site at 3050 m and as oc- curring primarily in single-species flocks. Here, we expand on Remsen’s (1985) data on diet, breeding condition, and body weights by including specimens from other localities in Bolivia and Peru. Of 11 stom- ach contents recorded, all consisted of ar- thropods. Of six specimens collected in Oc- tober, three were in breeding condition (males with testes 4 X 4 and 6 X 3 mm; female with brood patch); a female collect- ed in March may have been breeding (ova- ry 5 X 5 mm, largest ovum 1 X 1 mm). Fjeldsa & Krabbe (1990) collected a male with enlarged testes in January. Only one of the 13 specimens from July and August was possibly in breeding condition: a male with testes 3 X 2 mm. The body masses of five males with “no” or “light” fat were 14.5, 15, 16, 18, and 18 g; one with “‘mod- erate fat’’ weighed 22.5 g. The body masses of 12 females with “no” or “light” fat ranged from 12.4 to 19.2 g (mean 15.2); one with “‘moderate”’ fat weighed 16 g. Body or tail molt was recorded on the la- bels of 6 of 14 specimens collected in July and August, but none of the six specimens from October. Foraging observations of C. f. gravesi are difficult to obtain because this species is wary and remains on or near the ground in dense undergrowth. Etymology.—It is a pleasure to name this distinctive taxon for Gary R. Graves. Not only has he studied plumage variation in the genus (Graves 1980), but his synthetic an- alyses of geographic variation of Andean birds, which have included Cinnycerthia 1014 wrens (Graves 1985), have contributed greatly to knowledge of Andean birds. We previously proposed the English name ‘“‘Superciliated Wren” for C. fulva (Brumfield & Remsen 1996), a slight mod- ification of Hellmayr’s (1934) English name for this taxon, “‘Superciliated Brown Wren.”’ We overlooked, however, that Hell- mayr and everyone since then used “‘Su- perciliated Wren” for Thryothorus super- ciliaris. Unfortunately, virtually all Thry- othorus species have superciliaries, whereas the presence of a strong superciliary distin- guishes C. fulva from other Cinnycerthia. Nevertheless, we do not wish to change a long-established English name. Therefore, we propose “‘Fulvous Wren” as the English name for C. fulva. Acknowledgments We thank the following for specimen loans: David Willard and Scott M. Lanyon (Field Museum of Natural History), Fran- ¢ois Vuilleumier (American Museum of Natural History), Mark B. Robbins and Frank B. Gill (Academy of Natural Scienc- es, Philadelphia), D. Scott Wood and Ken- neth C. Parkes (Carnegie Museum of Nat- ural History), and Gene Hess (Delaware Museum of Natural History). Richard C. Banks, Frederick M. Bayer, and C. Brian Robbins provided valuable comments on the manuscript. We thank the following for permits and assistance with LSUMZ field- work in Peru and Bolivia in 1980 and 1981 that produced the specimens examined herein: the Academia Nacional de Ciencias (La Paz), Gast6n Bejarano, L. C. Binford, L. Campos L., A. P. Capparella, Direcci6n General Forestal (Ministerio de Agricultura, Lima), Groves Construction Company, Lin- da S. Hale, Tom and Jo Heindel, S. M. Lan- yon, Manuel Sanchez S., James Solomon, T. S. Schulenberg, A. Urbay T., and D. A. Wiedenfeld. Babette C. Odom, John S. Mcllhenny, E. W. Mudge, and H. Irving and Laura Schweppe sponsored LSUMZ PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON field-work in 1980 and 1981 in Peru and Bolivia. Literature Cited Brumfield, R. T., & J. V. Remsen, Jr. 1996. Geographic variation and species limits in Cinnycerthia wrens of the Andes.—Wilson Bulletin 108:205— DAT Fjeldsa, J., & N. Krabbe. 1990. Birds of the high An- des. Zoological Museum, University of Copen- hagen, Denmark, 876 pp. Graves, G. R. 1980. Relationship of white facial feath- ering to age and locality in Peruvian Cinnycer- thia peruana.—Bulletin of the British Ornithol- ogists’ Club 100:149—150. . 1985. Elevational correlates of speciation and intraspecific geographic variation in plumage of Andean forest birds.—Auk 102:556—579. Haffer, J. 1974. Avian speciation in tropical South America. Nuttall Ornithological Club Publica- tions, no. 14, 390 pp. Hellmayr, C. E. 1934. Catalogue of birds of the Amer- icas. Field Museum of Natural History Publi- cations, Zoological Series, Vol. 13, pt. 7, 531 PPp- Remsen, J. V., Jr. 1981. A new subspecies of Schi- zoeaca harterti with comments on taxonomy and natural history of Schizoeaca (Aves: Fur- nariidae).—Proceedings of the Biological So- ciety of Washington 94:1068—1075. . 1984. Geographic variation, zoogeography, and possible rapid evolution in some Crani- oleuca spinetails.—Wilson Bulletin 96:515— 5233). . 1985. Community organization and ecology of birds of high elevation humid forest of the Bolivian Andes.—Pp. 733-756 in P. A. Buckley et al., eds., Neotropical Ornithology. Ornitho- logical Monographs No. 36. . 1993. Zoogeography and geographic variation in Atlapetes rufinucha (Aves: Emberizinae), in- cluding a distinctive new subspecies, in south- ern Peru and Bolivia.—Proceedings of the Bi- ological Society of Washington 106:429—435. Ridgely, R. S., & G. Tudor. 1989. The birds of South America. Vol 1. The oscine passerines. Univer- sity of Texas Press, Austin, 516 pp. Ridgway, R. 1912. Color standards and color nomen- clature. Published by the author. Washington, D.C., 43 pp. Terborgh, J. 1971. Distribution on environmental gra- dients: theory and a preliminary interpretation of distributional patterns in the avifauna of the Cordillera Vilcabamba, Peru.—Ecology 52:23— 40. Vaurie, C., J. S. Weske, & J. W. Terborgh. 1972. Tax- onomy of Schizoeaca fuliginosa (Furnariidae), VOLUME 111, NUMBER 4 1015 with description of two new subspecies.—Bul- the Apurimac Valley of Peru with respect to en- letin of the British Ornithologists’ Club 92:142— vironmental gradients, habitats, and related spe- 144. cies. Unpublished Ph.D. dissertation, University Weske, J. S. 1972. The distribution of the avifauna in of Oklahoma, Norman. 137 pp. ce On awe . “up ead WAR “Sypeiaae Sat {Bini Ver atten POG, eakehe tio : a rah ri ae ae \osciay ae + pity ry! hee & Ga ete. aM a: ; Po 3 , ig 2) ae i —_ : 4 Sabee LP 8) ieee rh Pee’... TRACI SUrwet Ayhaee A Soaks ' were uk ts ee ~ yh, cd he it Cee cr em INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (botany, zoology, and paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers are published in English (except for Latin diagnoses/descriptions of plant taxa), with an Abstract in an alternate language when appropriate. Submission of manuscripts.—Submit three copies of each manuscript in the style of the Proceedings to the Editor, complete with tables, figure captions, and figures (plus originals of the illustrations). Mail directly to: Editor, Proceedings of the Biological Society of Wash- ington, National Museum of Natural History NHB-108, Smithsonian Institution, Washington, D.C. 20560. 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CONTENTS Sponges, genus Mycale (Poecilosclerida: Demospongiae: Porifera), from a Caribbean mangrove and comments on subgeneric classification Eduardo Hajdu and Klaus Ritzler First record and new species of Gastrophanella (Porifera: Demospongiae: Lithistida) from the central East Pacific Patricia Gomez Asexual reproduction in Linuche unguiculata (Swartz, 1788) (Scyphozoa: Coronatae) by plan- uloid formation through strobilation and segmentation Fabio Lang da Silveira and André Carrara Morandini Acteonina permiana, a new species from the Permian Coyote Butte Formation, central Oregon (Mollusca: Gastropoda: Actaeonidae) Rex Alan Hanger and Ellen E. Strong Two new species of Spionidae (Polychaeta) from Tahiti, French Polynesia Pat Hutchings, Patrick Frouin, and Christian Hily New species of bathyal and abyssal Sabellariidae (Annelida: Polychaeta) from near New Cal- edonia (southwest Pacific Ocean) Jean-Paul Lechapt and David W. Kirtley Two new species of Nereis (Polychaeta: Nereididae) from Todos Santos Bay, Ensenada, Baja California, México J. A. de Le6n-Gonzalez and V. Diaz-Castaneda Marphysa belli (Polychaeta: Eunicidae) and two related species, Marphysa oculata and M. totospinata, a new species, with notes on size-dependent features Hua Lu and Kristian Fauchald Pseudechiniscus asper, a new Tardigrada (Heterotardigrada: Echiniscidae) from Hokkaido, northern Japan Wataru Abe, Kazuo Utsugi, and Masatsune Takeda Parapetalophthalmus suluensis, a new genus and species (Crustacea: Mysidacea: Petalophthal- midae) from the Sulu Sea Masaaki Murano and Manuel Rafael Bravo Diastylis tongoyensis, a new diastylid (Crustacea: Cumacea) from the northern central coast of Chile, with an amendment to the description of Diastylis crenellata Watling & McCann, 1997 Sarah Gerken and Les Watling Redescription of the poorly known porcelain crab, Lissoporcellana nakasonei (Miyake, 1978) (Crustacea: Decapoda: Anomura: Porcellanidae) Masayuki Osawa A new genus and species of ghost shrimp (Crustacea: Decapoda: Callianassidae) from the Atlantic Ocean Richard W. Heard and Raymond B. Manning A new genus of ghost shrimp from Japan (Crustacea: Decapoda: Callianassidae) Raymond B. Manning and Akio Tamaki Lamoha hystrix, a new species of deep-water porter crab (Crustacea: Decapoda: Brachyura: Homolidae) the central Pacific Peter ie Nie A new freshwater crab of the genus Neostrengeria Pretzmann, 1965, from Colombia (Crusta- cea: Decapoda: Brachyura: Pseudothelphusidae), with a key to the species of the genus Martha R. Campos and Rafael Lemaitre A new species of mud shrimp, Upogebia toralae, from Veracruz, México (Decapoda: Thal- assinidea: Upogebiidae) Austin B. Williams and Jorge L. Hernandez-Aguilera Pinnotheres malaguena Garth, 1948, a new member of the genus Fabia Dana, 1851 (Crustacea: Brachyura: Pinnotheridae) Ernesto Campos and Raymond B. Manning Cave chaetognaths in the Canary Islands (Atlantic Ocean) F. Hernandez and S. Jiménez The discovery of Glyphocrangon stenolepis Chace (Decapoda: Caridea: Glyphocrangonidae) from Taiwan and Japan, with notes on individual variation Tomoyuki Komai, Tin-Yam Chan, and Ding-An Lee A new stomatopod (Crustacea: Malacostraca) of the genus Harpiosquilla Holthuis, 1964 from Taiwan and Australia Shane T. Ahyong, Tin-Yam Chan, and Y. J. Laio A new species of the genus Bellator (Pisces: Triglidae), with comments on the trigloids of the Galapagos Islands William J. Richards and John E. McCosker A new species of Polyipnus (Teleostei: Stomiiformes) from the western Indian Ocean, with comments on sternoptychid ecology Antony S. Harold, James H. Wessel, III, and Robert K. Johnson A review of western north Atlantic species of Bembrops, with descriptions of three new species, and additional comments on two eastern Atlantic species (Pisces: Percophidae) Bruce A. Thompson and Royal D. Suttkus A new species of cardinalfish (Apogonidae) from the Philippines, with comments on species of Apogon with six first dorsal spines Thomas H. Fraser The Neotropical catfish genus Epapterus Cope (Siluriformes: Auchenipteridae): a reappraisal Richard P. Vari and Carl J. Ferraris, Jr. Two new subspecies of Cinnycerthia fulva (Aves: Troglodytidae) from the southern Andes J. V. Remsen, Jr. and Robb T. 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