•» ouruiiSNi NVINOSHJ.IWS S3 I HVH 8 II LIBRARIES SMITHSONIAN m ^ 5 co ■ uj x^nn^x g* lli o - '-asss-' o BR ARIES SMITHSONIAN^INSTITUTION NOIJ.(UllSNl”WINOSHJLIlNS DimiiiSNi NviNOSHiiws^saiavaan li b rar i e s wsm ithson ian ~ i n st it 2L — |g Z' w- W 2 J 1 in O 2 5 ^y. pxx 5 BRARIEs‘OSMITHS0NIAN_INSTITUTI0N^N0linillSNI^NyiN0SHilWS^S3ia\ m '* JT| ^ BRAR I E S ^SMITHSON IAN INSTITUTION^ MOIinillSNI ^MVINOSHJLi^lS^ S3 I H\! 12 ^ z * co 2 co 2 XCtfWt^X < * S 2 2 O CO X n ■SS mniiiSNi ^ N^Aiiy NVIN0SH1IWS co x CO o s • > ‘ s S3IMVdan^LIBRARIESWSI X CO X CO o >* ITHSONIAN INSTITI i lq vi 1 1 noun inn mwimuiiwm ivuixiiaixqii! iv v i muonii oasavac co ^ z v cn z w ~ < a s _*«$& . < s § (1^ Jl?) § 8 (f£ ^$1 So d» I i wi^AiWi, "'IjjP’ > Xii***^ ^ >'y s >’ nilJLSNI^NVINOSHUWS^SB I dVHan\| BRAR I ES^SMITHSONIAN^fNSTITUTl ^ ^ . 5 w ^ 5 ^ ^ ^22^ oo ^OA^X yi w M'ASV}'>' O " ^ Q 2 -J z _ „ RARIES SMITHSONIAN INSTITUTION NOIlfUIJLSNI NVIN0SH1IINS $3ldVdS 2 r- _ z us 33 > 30 m ^ x^iiis^x m LniUSNI NVINOSHillAIS S3 I HVH 8 l“Tl IBRARI ES^SMITHSONIAN'lNSTITUT 5 > v- 2 ^ g RARIESWSMITHS0NIAN_INSTITUTI0NwN0M.nXllSNI__NVIN0SHilWSWS3iaVa! < |o^ '^jjNy c ^ l°l 2 O 2 6 ^ 2 .nillSNJ^NVINOSHimS S3 I d Vd 8 II LI B RAR I E $Z SMITHSONIAN^ I NSTI TUT z r- z r* 2 ° 33 #3- > [p 2 v m W' ^ xjvasv^x jn ^ jn - ' CO ^ 2 RAR I ES SMITHSONIAN INSTITUTION NOlifUIJLSM NVIN0SH1IINS S3f dVd* z • in z w ^ < V S < z o 00 § 2 ')^v >' 2 x^yosv^x > .fUiisNi_NviN0SHJ.iHs“s3 1 ava a nf li brar i es^smithsonian jnstituti S212^\ W X. - w ^T?ov5n “ Ab. cz w 2 ,ST r 5 £•” ' x' RARIES SMITHSONIAN INSTITUTION^NOlinillSNI JNVIN0SH1IWSZS3 I ava I 5 *"* . z r* z ^ S Xsj>s^ | \C||y s « ^ j; — ..... — CO PSYCHE A Journal of Entomology Volume 91 1984 Editorial Board Frank M. Carpenter, Editor H. W. Levi W. L. Brown, Jr. Alfred F. Newton, Jr. E. O. Wilson M. D. Bowers B. K. Holldobler J. M. Carpenter Published Quarterly by the Cambridge Entomological Club Editorial Office: Biological Laboratories 16 Divinity Avenue Cambridge, Massachusetts, U.S.A. The numbers of Psyche issued during the past year were mailed on the following dates: Vol. 90, no. 4, for 1983, April 6, 1984 Vol. 91, nos. 1-2, for 1984, November 15, 1984 ISSN 0033 2615 PSYCHE A JOURNAL OF ENTOMOLOGY founded in 1874 by the Cambridge Entomological Club Vol. 91 1984 No. 1-2 CONTENTS Adelpha (Nymphalidae): Deception on the wing. Annette Aiello 1 Earwigs of the California Channel Islands, with Notes on Other Species in California (Dermaptera) Scott E. Miller 47 Nesting biology and defensive behavior of Mischocyttarus (Monocyttarus) mexicanus cubicola (Vespidae: Polistinae). Henry R. Hermann and Jung- Tai Chao 51 Revision of the milvina group of the wolf spider genus Pardosa (Araneae: Lycosidae). C. D. Dondale and J. H. Redner 67 The identity of Chaetoclusia affinis Johnson and its placement in Sobarocephala Czerny (Diptera: Clusiidae). Norman £. Woodley . 119 The influence of microhabitat and prey availability on burrow establishment of young Geolycosa turricola (Treat) and G. micanopy Wallace (Araneae: Lycosidae): a laboratory study. G. L. Miller 123 Recently recognized types of some Homoptera described by Dr. Asa Fitch. Jeffrey K. Barnes 133 Production and use of secretions passed by males in pieris protodice (Lepidoptera, Pieridae). Ronald L. Rutowski 141 Female monogamy and male competition in Photinus collustrans (Coleoptera: Lampyridae) Steven R. Wing 153 Non-diapause overwintering by Pieris rapae (Lepidoptera: Pieridae) and Papilio zelicaon (Lepidoptera: Papilionidae) in California: adaptive- ness of type III diapause-induction curves. Arthur M. Shapiro 161 CAMBRIDGE ENTOMOLOGICAL CLUB Officers for 1983-1984 Margaret Thayer Donald S. Chandler Scott E. Miller Frank M. Carpenter Andrew Spielman Mary Hathaway EDITORIAL BOARD OF PSYCHE F. M. Carpenter, (Editor), Fisher Professor of Natural History, Emeritus, Harvard University W. L. Brown, Jr., Professor of Entomology, Cornell University and Associate in Entomology, Museum of Comparative Zoology B. K.. H0LLDOBLER, Professor of Biology, Harvard University H. W. LEVI, Alexander Agassiz Professor of Zoology, Harvard University M. D. BOWERS, Assistant Professor of Biology, Harvard University ALFRED F. Newton, Jr., Curatorial Associate in Entomology, Harvard University E. O. WILSON, Baird Professor of Science, Harvard University PSYCHE is published quarterly by the Cambridge Entomological Club, the issues appearing in March. June, September and December. Subscription price, per year, payable in advance: $13.00 domestic and foreign. Single copies. $4.00 Checks and remittances should be addresssed to Treasurer, Cambridge Entomological Club, 16 Divinity Avenue. Cambridge, Mass. 02138. Orders for missing numbers notices of change of address, etc., should be sent to the Editorial Office of Psyche. 16 Divinity Avenue, Cambridge. Mass. 02138. For previous volumes, see notice on inside back cover. IMPORTANT NOTICE TO CONTRIBUTORS Manuscripts intended for publication sho#ld be addressed to Professor F. M. Carpenter, Biological Laboratories, Harvard University, Cambridge, Mass. 02138. Authors are required to bear part of the printing costs, at the rate of $29.00 per printed page. The actual cost of pieparing cuts for all illustrations must be borne by contributors: the cost for full page plates from line drawings is ordinarily $10.00 each, and for full page half-tones, $12.00 each; smaller sizes in proportion. There is ordinarily no additional charge for setting tables of less then six columns; for tables of six or more columns the cost is $25 per page. President . . . p ... . Vice-President Secretary Treasurer Executive Committee Psyche, vol. 90, no. 4, for 1983, was mailed April 6, 1984 The Lexington Press, Inc., Lexington, Massachusetts PSYCHE Vol. 91 1984 No. 1-2 ADELPHA (NYMPH ALIDAE): DECEPTION ON THE WING* By Annette Aiello Smithsonian Tropical Research Institute P.O. Box 2072, Balboa, Republic of Panama Introduction For the past century, lepidopterists have puzzled over the genus Adelpha Hiibner, in an attempt to discover the secret character or combination of characters which might lead to a satisfying classification of the 100 or more butterfly species included in this large neotropical group. Several approaches have been tried (Godman & Salvin, 1884, 1901; Fruhstorfer, 1907; Forbes, un- published manuscript): wing pattern (both upperside and lower- side), wing venation, genitalia, and various combinations of these. If one attempts to coordinate all the information available, the result is a hopeless tangle. As a result, the most obvious set of characters (wing pattern) traditionally has been used in classification; other character groups (genitalia, larvae, pupae) which appear to confuse the situation have been largely ignored. A new strategy is needed, one which would both evaluate the reliability of the different groups of characters already surveyed, and search anew for overlooked sources of information. It was the purpose of my research to review what is known of the immature stages of Adelpha species and, based upon that information plus my own observations made in Panama between 1978 and the present, to make speculations regarding species relationships. 'Manuscript received by the editor January 12, 1984. 2 Psyche [Vol. 91 History of Classification The first attempt at classification within Adelpha was made by Godman & Salvin (1884, 1901) with their revision of the 32 species reported from Central America. At that time some 70 species were known for the genus. In their treatment, Godman & Salvin discuss Adelpha and its relationship to Limenitis and note that several species (bredowi, populi, Camilla, lorquini) are difficult to assign to either genus. Distribution of such characters as eye pubescence, venational differences, variation in proportions of male leg seg- ments, and peculiarities of male genitalia within Adelpha- Limenitis is surveyed. Classification of the 32 species begins with isolation of A. bredowi (eyes smooth in front) from all others (eyes hairy in front) and continues by arranging the other 31 species using gross features of wing pattern. The result is eleven groups, six of which are represented by single species. With Fruhstorfer (1907) came the first and only published revision of the entire genus Adelpha. The 90 species treated are assigned to two main groups based upon the length of the forewing discal cell; short = Adelpha group, elongate = Heterochroa group. Upon this division, Fruhstorfer comments, “Anatomically there are also two series [male genitalia with or without clunicula] of species distinguishable. They, however, do not agree with those based upon the structure [discal cell length].” The linear arrangement of species within the largest group ( Heterochroa with 82 species) reflects presumed relationships based upon wing pattern features. Fruhstorfer touches upon but does not pursue the possibility of a closer than realized alliance between New and Old World groups. In his introduction he notes that the male clasping organs of Adelpha “. . .approach those of the [Old World] genus Pantoporia (Athyma) in such a way that. . .it would be quite impossible to ascertain where organs or photes [sic] of them belong to, which are not denomi- nated.” Concerning the male valves, he further observes, “. . .there exist also nearly square ones with 2 or 3 small acicular teeth (resembling a Limenitis [Mo duza] pro cr is from India and Ceylon).” Fruhstorfer’s revision included a number of misidentifications, which presumably were corrected by Hall (1938) following examin- ation of the original material. At the time of his death (1968) W. T. M. Forbes had made a good deal of progress on a revision of Adelpha, and his manuscript is in 1984] Aiello — Genus Adelpha 3 the Archives of the Museum of Comparative Zoology, Harvard University. About half of this manuscript is in a nearly illegible hand; half is typewritten. The typed portion includes part of an introduction, notes on each species, a key (based upon wing pattern) to species and species groups, and descriptions of four new species. Included also are more than 70 genitalic illustrations (inside of right valve) by Howarth. Forbes had analyzed these latter and was in the process of constructing a key to genitalia. His two approaches, of wing pattern and genitalia, yielded different species groupings, and it is not clear whether he favored one of these, or intended to use a combination of the two in his final classification. Of the immature stages of Adelpha, very little has been said, and what has been said has been largely ignored for the simple reason that this new information appears to confuse matters rather than clarify them.1 Forbes (uncompleted manuscript) had read Moss’s (1933) paper on Adelpha larvae and pupae, and had examined several pupal skins of Adelpha and Old World Limenitini in the collections of the British Museum (Natural History) when he commented, “The larvae and pupae are highly varied and unless a high percent are misdetermined, show characters wholly incongru- ent with adult structures and patterns.” The fact that, within Adelpha, the study of adult characters results in species-alignments different from those obtained by consideration of the immature stages and/or genitalia, suggests that at least one set of characters is unreliable or perhaps even deliberately deceptive. Immature Stages Due to taxonomic confusion within Adelpha, it is impossible to know how many species actually are represented by the 34 or so life history accounts published for this genus. My estimate is that at least 24 species are illustrated (including illustrations in this paper) as final instar larva, or pupa, or both, many of these by more than one author. Most accounts, whether illustrated or not, include a foodplant record. 'This author has located only four publications (Moss, 1933; Muller, 1886; Young, 1974; Comstock & Vasquez, 1960) which figure any immature stages of Adelpha; several publications present descriptions only. 4 Psyche [Vol. 91 In the present paper, all illustrated reports are reviewed to see whether it is possible to distinguish natural species groups within the genus Adelpha. Oviposition Judging from the few scattered reports (Table 1) of oviposition in the Limenitidini, eggs are laid singly and are usually placed at the tips of leaves. While many accounts do not specify whether placement is on the upper or lower surface, the majority that do specify, report upper surface oviposition. Adelpha iphicla females alight upon a leaf and walk backwards while searching for the desired oviposition site with the tip of the abdomen; the egg is then placed and the butterfly takes flight before her next oviposition, which may be on the same leaf. In the case of lower surface oviposition, the female merely bends her abdomen around the edge of the leaf and touches its tip to the leaf. Among the six species of Adelpha whose eggs were collected in Panama, four (Table 1) place their eggs along damaged portions of leaves, especially on jagged points, as well as at the leaf tip, and one of these (A. iphicla) is about as likely to lay eggs along an intact leaf margin as at the tip. As well, a female A. iphicla may return to the same leaf several times and place as many as four eggs on one leaf, often on its undersurface. Eggs Eggs of the seven Adelpha species ( basiloides , cocala, cytherea, marcia, iphicla, melanthe, phylaca aethalia, and salmoneus) exam- ined by this author, were all similarly sculptured (figure 1) with pits (hexagonal due to packing) and seta-like projections (one from each junction of three pits). This same sculpture type is figured for Limenitis by both 2Scudder (1889) and Eltringham (1923). Young’s (1974) description of the egg of Adelpha leucophthalma as having the seta-like projections “aris[ing] from the facets,” is doubtful. Also doubtful is the account by Comstock and Vazquez (1960) which describes and illustrates the egg of A. celerio as having convex rather then concave hexagons. Such an error is easily made, as these 2 As Basilar chia. 1984] Aiello — Genus Adelpha 5 Table 1. Placement of egg on leaf by various members of the Limenitidini (Nymphalidae) BUTTERFLY SPECIES EGG PLACE- MENT REFERENCE Adelpha basiloides Bates Ti, Di Aiello Adelpha celerio diademata Fruh. T Comstock & Vazquez Adelpha cocala Cr. Ti, (Dl) (1960, pg. 407) Aiello Adelpha cytherea (?) t MUller (1886, pg. 484) Adelpha iphicla L. Tl(t), Mi(t) Aiello Adelpha iphicla t MUller (1886, pg. 484) Adelpha isis Dru. Ti, Di MUller (1886, pg. 481) Adelpha leucophthalma tegeata Fruh. Di Young (1974, fig. 2) Adelpha melanthe Bates Di Aiello Adelpha phylaca aethalia Feld. Ti Aiello Adelpha plesaure Ti MUller (1886, pg. 484) Adelpha salmoneus Butl. Ti, (Di) Aiello Adelpha serpa Ti MUller (1886, pg. 484) Adelpha syma Godt. T Hoffmann (1937, pg. 212) Athyma nefte Cr. (as Parathyma nefte Cr.) Ti, (Ml) Morrell (1954, pg. 160) Athyma opalina Kollar t Robson (1894, pg. 338) Lasippa tiga Moore (as Neptis heliodore Fruh.) T Morrell (1954, pg. 160) Limenitis archippus (as Basilar chia archippus) Tt Scudder (1889, fig. 16) Limenitis lorquini burrisonii Maynard T Dornfeld (1980, pg. 61) Moduza procris Cr. T Morrell (1954, pg. 157) Neptis nata Moore T Morrell (1954, pg. 162) T = leaf tip (tooth, in case of A syma), M = leaf margin (intact), D = damaged portion of leaf, i = upper surface of leaf, t = lower surface of leaf, ( ) = less commonly Figure 1. Eggs of Adelpha (left) and Doxocopa (right). 6 Psyche [Vol. 91 eggs can present an optical illusion, however, an edge view of a partly eaten egg shell shows clearly that the hexagons are indeed pits. Comstock and Vazquez (1960) also describe and figure a differently sculptured egg, which they call A. iphicla with the reservation that it might actually be an egg of 3 Doxocopa. Because their larvae died soon after hatching, the authors never knew that they had indeed been fooled by the ovipositing Doxocopa female, a mimic of Adelpha. My own rearing of Doxocopa laure (LOT 83-6) was from two eggs with pattern (figure 1) identical to that in figure 33 of their paper. My reared Adelpha iphicla were from eggs patterned as those of all the other Adelpha species examined so far. Possibly this egg sculpture pattern will be found throughout the Limenitini and may prove useful in defining tribal limits. Development Time Oviposition was observed only for Adelpha iphicla, and each of the three fresh eggs collected required 5 days development before the first instars emerged. Of five other species collected as eggs, of unknown age, three had longer minimum egg-development times than did A. iphicla: A. basiloides (LOT 82-65), one egg hatched after 6 days; A. cytherea (LOT 83-3), one egg hatched after 7 days; and A. salmoneus (LOT 83-14), two eggs hatched after 6 days. Because individuals were collected at various stages of devel- opment, total time from hatching to eclosion could be determined for only a few individuals of ten species. In spite of the fact that the resulting data (Table 2) is not uniform, it is clear that development time is variable, and often more so within a species than between species. The extreme example of wide variation is in A. basiloides which has a longer development time than other species, and which sometimes passes through six instars instead of the usual five. Five 6-instar individuals of A. basiloides were encountered, and these were from four separate rearing lots, each of which included 5-instar individuals as well. Each of these lots was collected on Amaioua corymbosa; larvae collected on Alibertia edulis and Bertiera guianensis did not produce extra instars. In spite of the additional instar, 6-instar individuals did not require longer development times 3 As Chlorripe. 1984] Aiello — Genus Adelpha 7 than 5-instar individuals, however, larvae on Amaioua, regardless of number of instars, were slower to develop (41-48 days) than were larvae on Alibertia or Bertiera (32-44 days). Larvae Miles Moss reared twelve species of Adelpha from Par&, Brasil, and illustrated (1933) the final instar larva and the pupa for each of them. In spite of the lack of detail in his illustrations and descriptions, and the fact that his scheme for numbering body segments omits abdominal segment-9, his little paper remains the masterpiece on Adelpha immatures. It was Moss who realized (page 15) that: . .systematists,. . .by a careful examination of certain hither- to unsuspected points of likeness or dissimilarity between the species in their early stages, may perhaps be led to modify the existing order and grouping of the butterflies of this difficult genus. It is just possible that a few unexpectedly close relationships may thus be established, while others at present confused, or regarded as near of kin, may be found to be more distantly related than was supposed.” He concluded that A. cytherea, and pseudococala are closely related, also delphicola and mesentina, and serpa and para'ena.. My analysis of published illustrations and live material lends support to Moss’s groupings and adds several more. FIRST INSTAR LARVAE First instar larvae, of all species which I have seen, appear identical in form, and are some shade of brown or grey. The head bears setae, but none of the4chalazae of later instars; pale bumps on the body are found where the future 4scoli will be, and the body is covered with tiny pale spots each centered with a minute seta. After their first meal, larvae take on the green color of their foodplant, although the head remains brown. SECOND INSTAR LARVAE From the second instar on, the head is ornamented with chalazae which give it a spiny appearance. The face has numerous round, Table 2. Duration (in days) of larval instars and pupal stage for twelve Adelpha species. Instar Total Time 8 Psyche [Vol. 91 justina m4 4 3, 4 3, 4 4-6 - 6-9 m30 Table 2. (Continued) Duration (in days) of larval instars and pupal stage for twelve Adelpha species. 1984] Aiello — Genus Adelpha 9 () = commonest number of days in cases of strong bias towards one end of range; otherwise, ranges are normal curves, m = minimum number of days in cases where full time for an instar is not known. 10 Psyche [Vol. 91 flatbottomed pits which first appear in small numbers in instar two, three, or four depending upon the species. These may be the same color as the rest of the head capsule or may be a contrasting color. Facial stripes appear in the fourth instar of some species as well. The body now bears stubby scoli (each with 3-5 radiating apical spines). These scoli are arranged in three rows (subdorsal, supra- spiracular, and subspiracular), thus, each body segment has three pairs of scoli. The subdorsal scoli on thoracic segments 2 & 3 and abdominal segments 2, 7, & 8 are very slightly larger than the others, but all are similar in form. Beginning with the second instar there are color and pattern changes in some species, but most are fairly uniformly colored (brown, green, or black) and have paler scoli and tiny spots, much as the first instar. In Adelpha basiloides, the larval color depends upon the foodplant; larvae are light brown or reddish brown on Amaioua, and dark brown on Alibertia and Bertiera. THIRD AND FOURTH INSTAR LARVAE From instar three on, the face is framed by two distinct rings of chalazae, and body scoli show further development, especially those scoli which will be the largest or most distinctive in the final instar. By late fourth instar, body scoli have become swollen at the bases due to developing final instar scoli inside. In these two instars, the scolus spines are pale, and as before, the body is speckled with tiny pale spots. A dorsal, paler patch (“saddle”) appears in the third or fourth instar of many species. In A. salmoneus and cytherea the “saddle” is faint and extends from abdominal segment 2 or 3 through segment 6. A. cocala and leucophthalma may have a “saddle” on abdominal segments 5-6. A. justina is pale dorsally from thoracic segment 2 through abdominal segment 8. A. basiloides, nr. paraena, and celerio have a triangular “saddle” with its base on the posterior portion of abdominal segment 4, and its apex at mid abdominal segment 6. In the latter two species the “saddle” is poorly defined; in A. basiloides it is sharply demarcated. “♦Terminology of Peterson (1962). chalazae(AE): a distinctly elevated cone-shaped area, bearing 1-3 simple setae; (SCOLUS(i): an elongated projection, bearing 4 or more setae or spines. 1984] Aiello — Genus Adelpha 11 FINAL INSTAR LARVAE Final instar larvae (Figures 4 & 5) of all species studied have several things in common. The head (Figure 2) has a spiny appearance owing to the numerous chalazae which frame the face. These seem to be fairly constant in number and position (Figure 3) and to vary from species to species mainly in their relative size. A. iphicla for example has a relatively smooth face because it lacks several of the chalazae found in other species. A. phylaca and especially melanthe are the spiniest in appearance due to numerous additional setae. Color and pattern also vary: A. celerio has a striped face, A. basiloides and A. cocala patterned ones. The pits of A. cytherea are darker than the rest of the face; the head of A. salmoneus is reddish and constrasts strongly with its green body. Each body segment bears three pairs of scoli (subdorsal, supra- spiracular, and subspiracular); there are no dorsal scoli. While the scoli are variable in form and length, in all species studied, those of the prothorax and abdomen- 1 are either very short or are reduced to a few spines. Usually the longest are the subdorsal scoli of meso- and meta-thorax and abdomen-2, -7 & -8, and the supraspiracular scoli of the meso-thorax. In many species the subdorsal scoli of abdomen-2 are the most distinctive in form and often are curved backwards. Body scoli are diverse but the various forms can be grouped into two main types: those which are terete (round in cross-section), and those which are flattened. The terete scolus, in its simplest form, is a short stalk with 3-5 spines radiating star-like from its apex (e.g., 5scoli A3-6 of A. celerio, cytherea, salmoneus, and justina ). More complex scoli are slender and longer with spines at intervals along their full length, and either with one to a few ascending spines towards the apex (thoracic scoli of many species, e.g., A. phylaca, melanthe, cocala, basiloides, cytherea, justina, salmoneus ), or with 3-5 radiating apical spines (e.g., scoli of A3-6 in A. phylaca, melanthe). Scoli also may be short and thick with a dense covering of spines (e.g., A2 of A. cocala, leucophthalma, basiloides ), or be club-shaped and more sparsely spined (e.g., A8 of basiloides). throughout the remainder of this paper, “scolus(i)” refers to the subdorsal set unless otherwise noted; T = thorax, thoracic segment; A = abdomen, abdominal segment. 12 Psyche [Vol. 91 Figure 2. Faces of final instar larvae of Adelpha species reared in Panam&: BAS = basiloides, CEL = celerio, COC = cocala, CYT = cytherea marcia, IPH = iphicla, MEL = melanthe, PHY = phylaca aethalia, SAL = salmoneus. 1984] Aiello — Genus Adelpha 13 Figure 3. Location of head chalazae 1-5, referred to in text. Flattened scoli have a plumose or leaf-like appearance (e.g., A. celerio, serpa, paraena ) which is due not only to the arrangement of spines in two opposite rows along the scolus, but also to the flattening and widening of those spines which may take on spatulate, elliptical, or lanceolate forms, and may be so crowded that they overlap one another. In the case of extremely condensed scoli, the spines appear to fuse with one another (see A. serpa in Muller, 1886). Larval Behavior Upon hatching, a larva eats some or all of its egg shell and then feeds on the leaf tip leaving the midrib intact. Larvae rest out on the midribs which they have exposed, and eventually extend them by addition of fecal pellets held in place with silk. Most larvae fashion several such supports during each instar, often using lateral veins instead of the midrib. When not feeding, Adelpha larvae rest out on their supports, facing away from the leaf; molting takes place on the support as well. After they molt to the final instar, larvae abandon their supports and thereafter rest on the upper surface of a leaf. 14 Psyche [Vol. 91 In addition to the above behavior, typical of many nymphalid butterfly species, first through fourth instar Adelpha larvae engage in an odd practice not known for other New World butterflies; they accumulate their fecal pellets, fastening them in place with silk, to form a mass which either surrounds the base of the support or is suspended beneath it. This mass may include bits of leaf in A. iphicla, or consist of more leaf than feces in A. phylaca and melanthe. The work of A. basiloides is the most distinctive because, in addition to the mass just described, this species consistently constructs a small, usually curved, larva-form mass on the leaf surface, several mm away from either the leaf edge or the usual mass. Upon viewing this artistry, one cannot help imagining that it serves as a decoy larva to discourage would-be predators. A single reared individual of A. iphicla also engaged in this behavior, but excepting that, I have observed it only in A. basiloides. Resting larvae position their bodies in one of several ways: (1) Straight Position (figures 4 & 5). Larvae in any instar may use this position, and typically when out on their supports they rest this way. (2) Front-Curved Position, as shown in Young (1974, fig. 2. B, C). In this position, the larva grasps its support with the prolegs only, and raises and curves its anterior portion (head through A2) so that the head is somewhat inclined, and the toracic scoli are directed forwards. Larvae about to molt use this position; the second instar in Young’s (1974, fig. 2C) photograph has a swollen pro-thorax and is probably preparing to molt to third instar. Final instar larvae of A.justina use a raised but uncurved version of this position instead of the usual final instar stance, described next. (3) Front-Arched-Rear-Up Position. This position is typical of final instar larvae but occurs in earlier instars as well. Involved are the raising and arching of the anterior portion of the body (head through A2), plus elevation of the posterior portion (A7-10). In addition, the thoracic scoli are directed forwards, and those of A2 are held backwards. In this position, the face and the area at the top of the arch (T3 through A2) are parallel to each other and to the substrate. This and the next position are assumed when the larva is disturbed. (4) Curled Position. The larva curls to one side into a “C” or “J” shape on the upper surface of the leaf; the rear portion (A7-10) may or may not be elevated. A. celerio often rests in this position, 1984] Aiello — Genus Adelpha 15 and when it does, the scoli on the outside of the curve stick out all around and the animal resembles a bit of moss. Moss (1933) reported the same behavior and appearance for A. thesprotia. Foodplants At first inspection, it would appear that larvae of Adelpha butterflies are not very particular about their foodplant selection. Indeed, some 56 plant species, representing 42 genera and 16 families have been reported as larval foodplants of Adelpha. Moreover, many Adelpha species are known to be polyphagous: A. melanthe, delphicola, and isis have each been reared on three plant genera, A. celerio on four, and A. cocala and iphicla each on eight. Cecropia (Cecropiaceae), Sabicea (Rubiaceae), and Vitex (Verben- aceae) are each attacked by five different species of Adelpha. However, when larval foodplants are grouped by the butterfly species which feed upon them (Table 3), a pattern does emerge: Adelpha species fall nicely into two main feeding groups: (1) Rubiaceous feeders, and (2) Non-rubiaceous feeders. With the following three exceptions, butterfly species that feed upon members of the Rubiaceae have not been reported on plants outside that family. A. boreas tizona was reared on both Rubiaceae and Ericaceae (Marquis and DeVries, unpublished); A. cocala, reared on seven members of the Rubiaceae, also has been reported on Emmotum of the Icacinaceae (Moss, 1933); A. syma, on Rubiaceae and Rosaceae. The record of A. iphicla iphicleola on Celtis (Ulmaceae) (Comstock and Vazquez, 1960) is in error; their butterfly was actually Doxocopa, misidentified as Adelpha. It is interesting to note that the rubiaceous genera, utilized as foodplants by Adelpha, belong to at least seven of the 18 tribes outlined for the Rubiaceae by Kirkbride (1982). Several non-rubiaceous feeders do show wide foodplant pref- erences, the extreme examples being: A. melanthe which feeds on three plant genera representing three families, and A. celerio which feeds on five genera in four families. The only common bond among the foodplants of these two butterflies seems to be that all are scabrous- or pubescent-leaved, second-growth trees. Some plants attract a more specialized group of feeders: Vitex (Verbenaceae) is attacked by A. abia, calliphane, epizygis, and jordani, which have not been reported on any other plants. A. naxia ipiphilca, also 16 Psyche [Vol. 91 Figure 4. Final instar larvae of Adelpha species reared in Panam&: abbreviations are the same as in Figure 2 except that CEL = nr. CEL; in addition nr. PAR = near paraena. 1984] Aiello — Genus Adelpha 17 Figure 4. (Continued) 18 Psyche [Vol. 91 Table 3. Larval foodplants reported for Adelpha species, arranged alphabetically by butterfly species. Adelpha species / FOODPLANT Plant Family Locality Reference ABIA Vitex sp. Verbenaceae Brasil MUller (1886) BASILEA Calycophyllum sp. Rubiaceae Cuba Dewitz (1879) BASILOIDES Alibertia edulis Rubiaceae Panama Aiello LOTS 79-121, 80- 38, 81-17, 81-42, 81- 49, 83-16, 83-42 Amaioua corymbosa Rubiaceae Panama Aiello LOTS 82-59, 82- 65, 82-70, 82-72, 83- 42 Bertiera guianensis Rubiaceae Panama Aiello LOTS 82-64, 82-73, 83-87 Ixora nicaraguensis Rubiaceae Panama Mallet in DeVries (unpub) BOETIA OBERTHURI Luehea seemannii Tiliaceae Costa Rica Mallet in DeVries (unpub) BOREAS TIZONA Sat y ria sp. Ericaceae Costa Rica Marquis in DeVries (unpub) Chomelia bispinosa Rubiaceae Costa Rica DeVries (unpub) BREDOWI CALIFORNIA Quercus chrysolepis Fagaceae California Orsak (1977) Quercus spp. (especially chrysolepis Fagaceae California Howe (1975) Quercus spp. Fagaceae California Dornfeld (1980) BREDOWI EULALIA Quercus spp. Fagaceae Mexico, US Ferris & Brown (1980) CALLIPHANE Vitex montevidensis Verbenaceae Brasil cited in Lima (1968) CALLIPHICLEA Ilex paraguariensis Aquifoliaceae Paraguay Jorgensen (1921) CELERIO Cecropia peltata Cecropiaceae Panama Coley LOT 15 Miconia argentea Melasto- Costa Rica Mallet in DeVries mataceae (unpub) Ochroma pyramidale Bombacaceae Panama Aiello LOT 82-41 Ochroma pyramidale Bombacaceae Panama Coley LOT 22 Urera sp. Urticaceae Costa Rica DeVries (unpub) 1984] Aiello — Genus Adelpha 19 Table 3. (continued) Adelpha species / FOODPLANT Plant Family Locality Reference CELERIO DIADEMATA Conostegia xalapensis Melasto- mataceae Mexico Comstock & Vazquez (1960) Miconia sp. Melasto- mataceae Mexico Comstock & Vazquez (1960) nr. CELERIO Heliocarpus popayanensis Tiliaceae Panama Aiello LOTS 82-75, 83-68 COCALA Calycophyllum candidissimum Rubiaceae Panama Aiello LOT 81-44 Calycophyllum candidissimum Rubiaceae Costa Rica Janzen 81-SRNP-740 in DeVries (unpub) Chomelia psilocarpa Rubiaceae Panama Aiello LOT 81-14 Emmotum nitensl Icacinaceae Brasil Moss (1933) Malanea sp. Rubiaceae Brasil Moss (1933) Pentagonia macrophylla Rubiaceae Costa Rica DeVries (unpub) Psychotria sp. Rubiaceae Costa Rica Marquis in DeVries (unpub) Uncaria tomentosa Rubiaceae Panama Aiello LOT 81-54 Warscewiczia coccinea Rubiaceae Panama Aiello LOT 82-66, 82-71, 82-74 CRETON Quercusl Fagaceae Mexico Miller & Miller (1970) CYTHEREA Sabicea aspera Rubiaceae Brasil Moss (1933) CYTHEREA MARCIA Sabicea panamensis Rubiaceae Panama Aiello LOT 83-129 Sabicea villosa Rubiaceae Panama Aiello LOTS 82-26 83-3 Sabicea villosa Rubiaceae Costa Rica DeVries (unpub) DELPHICOLA Bombax munguba Bombacaceae Brasil Moss (1933) Cecropia sp. Cecropiaceae Brasil Moss (1933) Pourouma sp. Cecropiaceae Brasil Moss (1933) DEMIALBA Rondeletia sp. Rubiaceae Costa Rica Haber in DeVries (unpub) DONYSA Quercusl Fagaceae Mexico Miller & Miller (1970) 20 Psyche [Vol. 91 Table 3. (continued) Adelpha species/ Plant FOODPLANT FAMILY LOCALITY REFERENCE EPIZYGIS Vitex montevidensis Verbenaceae EROTIA Tetrapteris sp. Malpighi- aceae FESSONIA Randia echinocarpa Rubiaceae Randia learstenii Rubiaceae IPHICLA Antirrhoea trichant ha Rubiaceae Alseis blackiana Rubiaceae Bathysa sp. nr. barbinervis Calycophyllum Rubiaceae candidissimum Rubiaceae Calycophyllum candidissimum Rubiaceae Gonzalea spicata Rubiaceae Gonzalea spicata Rubiaceae Isertia haenkeana Rubiaceae Rondeletia panamensis Rubiaceae Uncaria (as Ourouparia) guianensis Rubiaceae Uncaria tomentosa Rubiaceae iphicla ephesa (as ephicla ephesa ) Bathysa sp. IPHICLA IPHICLEOLA Calycophyllum Rubiaceae candidissimum Rubiaceae Brasil cited in Lima (1968) Brasil Milller (1886) Costa Rica Costa Rica Janzen 79-SRNP-216 in DeVries (unpub) Janzen 79-SRNP-725 in DeVries (unpub) Panama Panama Aiello LOTS 81-25, 82-27, 83-41 Aiello LOTS 82-36, 82-56, 82-63, 82-69 Brasil MUller (1886) Panama Aiello LOTS 81-34, 81-78 Cuba W.I. W.I. Panama Panama Riley (1975) Barcant (1970) Riley (1975) Aiello LOT 81-16 Aiello LOTS 83-101, 83-111 Brasil Panama Moss (1933) Aiello LOTS 81-46, 81-51 Brasil cited in Lima (1968) Costa Rica Janzen 79-SRNP-135 in DeVries (unpub) iphicla iphicleola ( Doxocopa misidentified as Adelpha ) Celtis sp. Ulmaceae Mexico Comstock & Vazquez (1960) 1984] Aiello — Genus Adelpha 21 Table 3. (continued) Adelpha species/ FOODPLANT Plant Family Locality Reference ISIS Cecropia pachystachia Cecropiaceae Brasil cited in Lima (1968) Cecropia pachystachia Cecropiaceae Brasil Muller (1886) Coussapoa schottii Cecropiaceae Brasil cited in Lima (1968) Coussapoa schottii Cecropiaceae Brasil M tiller (1886) Pourouma acutiflora Cecropiaceae Brasil cited in Lima (1968) Pourouma acutiflora Cecropiaceae Brasil M tiller (1886) JORDANI Vitex triflora Verbenaceae Brasil Moss (1933) JUSTINA JUSTINA Alternate-leaved plant ? Panama Aiello 83-23, 83-67 LARA Cecropiaceae Trinidad M. J. W. Cock (pers. comm.) LEUCOPHTHALMA Pentagonia wendlandia Rubiaceae Costa Rica Young (1974) Undetermined Rubiaceae Panama Aiello LOTS 83-22, 83-24, 83-25 Undetermined Rubiaceae Costa Rica DeVries (unpub) MELANTHE Cecropia sp. Cecropiaceae Costa Rica DeVries (unpub) Trema micrantha Ulmaceae Panama Aiello LOT 83-8 Trema micrantha Ulmaceae Costa Rica DeVries (unpub) Ur era sp. Urticaceae Costa Rica DeVries (unpub) MELONA Malanea sp. Rubiaceae Brasil Moss (1933) MESENTINA Pourouma sp. Cecropiaceae Brasil Moss (1933) MYTHRA Bathysa sp. nr. barbinervis Rubiaceae Brasil Mtiller (1886) NAXIA IPIPHILCA Piper areanum Piperaceae Costa Rica Marquis in DeVries (unpub) Vitex cooperi Verbenaceae Costa Rica DeVries (unpub) paraEna Isertia longiflora Rubiaceae Brasil Moss (1933) Remijia amazonica Rubiaceae Brasil Moss (1933) 22 Psyche [Vol. 91 Table 3. (continued) Adelpha species/ FOODPLANT Plant Family Locality Reference nr. paraEna Combretum decandrum Combreta- Panama Aiello LOT 82-55 ceae PHLIASSA Alibertia edulis Rubiaceae Brasil Moss (1933) Bertiera guianensis Rubiaceae Brasil Moss (1933) PHYLACA AETHALIA Cecropia obtusifolia Cecropiaceae Panama Aiello LOTS 82-39, 82-68, 82-76, 83-78 Cecropia peltata Cecropiaceae Panama Aiello LOT 81-70 Cecropia peltata Cecropiaceae Panama Coley LOT 5 Cecropia sp. Cecropiaceae Panama Aiello LOT 82-40 PLESAURE Bathysa ? sp. Rubiaceae Brasil Muller (1886) PSEUDOCOCALA Sabicea aspera Rubiaceae Brasil Moss (1933) pseudococala? (as nr. cocala) Sabicea sp. Rubiaceae Brasil Muller (1886) SALMONEUS Sabicea panamensis Rubiaceae Panama Aiello LOTS 82-37, 83-14 SERPA Miconia minutiflora Melasto- Brasil Moss (1933) mataceae Misc. spp. Melasto- Brasil MUller (1886) mataceae SERPA HYAS Ilex paraguariensis Aquifoliaceae Barsil cited in Lima (1968) sophax (= zalmona zalmona) SYMA Rubus fructicosus Rosaceae Brasil Muller (1886) Rubusl sp. (“bramble”) Rosaceae Brasil Jones & Moore (1882-3) Cephalanthus sarandi Rubiaceae Brasil cited in Lima (1968) THESPROTIA? Alternate-leaved climber ? Brasil Moss (1933) TRACTA Undetermined Rubiaceae Costa Rica Haber, Chacon in DeVries (unpub) ZALMONA ZALMONA (as SOphax) Sabiceae aspera Rubiaceae Costa Rica DeVries (unpub) 1984] Aiello — Genus Adelpha 23 reared on Vitex, has been found on only one other plant species {Piper, Piperaceae). Pupa Adelpha pupae (Figures 5 & 6) are diverse in form, yet have several features in common: (1) The body is slender and tapers towards the cremaster, so that the posterior wing margins protrude and appear as keels. (2) Segments T2 and A2 are expanded dorsally to form two projections of varying shape and size. The thoracic projection is the smaller of the two and may be pointed and directed posteriorly (A. salmoneus, cytherea, iphicla, justina ), or pointed and oriented almost perpendicularly to the body (A. melanthe, phylaca aethalia ), or it may be reduced to a slight hump (A. celerio ), or smooth curve {A. basiloides ). The abdominal projection is the larger and more variable of the two. It may be short, pointed, and directed anteriorly {A. basiloides, justina ), a bit larger and curved anteriorly {A. iphicla, celerio, cytherea, coca la), even larger and directed posteriorly {A. sal- moneus), or it may be a huge laterally flattened hook {A. melanthe, phylaca aethalia). (3) The head usually has a pair of apical projections (“horns”) of varying shape and size. These may be long, slender, and slightly curved {A. celerio), sickle-shaped and recurved to the sides (A. Figure 5. Final instar larva and pupa of Adelpha justina justina, (JUS) reared in Panama. 24 Psyche [Vol. 91 Figure 6. Pupae of Adelpha species reared in Panama: abbreviations are the same as in Figures 2 & 4; CEL = celario. 1984] Aiello — Genus Adelpha 25 basiloides ), shaped like tiny asymmetrical leaves ( A . coeala, leucophtha/ma), small and triangular, like cat ears (A. iphicla, cytherea, salmoneus), smaller triangles which are bent to the sides (A. justina, phylaea aethalia), or they may be reduced to two tiny rounded projections (A. melanthe). (4) Pupal color varies from pearly white, through straw-color, brown, green, copper, or shimmering gold or silver. Whatever its color, an Adelpha pupa gives the impressing that it is empty or diseased. The pearly white pupa of A. basiloides is especially deceptive; it appears more to be an abandoned skin than a solid object. The shimmering silver pupae of A. celerio and nr. paraena have black sutures and both give the impression that they are transparent with black lines showing through from the other side. As far as I know, the only other instance of a totally silver pupa occurs in Mechanitis of the Ithomiidae. The pupa of A. salmoneus is coppery brown and resembles a dead or diseased individual. Adelpha Species Groups In exploring for possible correlations between pupal and larval types, I began with species whose pupae have a large flat hook-like projection from the dorsum of A2. As it turned out, all have similar larvae, foodplants, and adult male genitalia, yet, the adult wing patterns are diverse. Further analysis revealed several other species groups that show similar correlations. Based upon this work, I have come to the conclusion that the genus Adelpha comprises five to seven natural groups, and possibly more. The only set of characters that doesn’t hold together within these groups is adult wing pattern. Some twenty species, for which adequate illustrations are available, are included in the classification which follows, and several others whose immatures were described only, are discussed. The outline and discussion of Adelpha species groups includes brief final instar descriptions for species reared by me in Panama, as well as comments upon previously published accounts of other species belonging to these groups. The major characteristics listed at the beginning of each group do not function as a key, rather, they represent defining characters for those groups, except as noted in parentheses, foodplants may host more than one butterfly group: members of groups I and II have been reared on plants from 26 Psyche [Vol. 91 a number of families, group III on Verbenaceae, and the remaining groups (IV-VII) on Rubiaceae. At this stage of our knowledge, an attempt to work out a phylogeny within Adelpha would be premature indeed; more information on life histories of Adelpha and its relatives (especially Old World genera) would be necessary to such an undertaking. ♦Key to Adelpha Species Groups Based upon Final Instar Larvae, Pupae, and Foodplants A. larva: body scoli, especially scolus of A2, flattened and feather-like; face with dark stripes. PUPA: shimmering silver with slender, very slightly twisted horns. FOODPLANTS: non-rubiaceous. (A. celerio, sp. nr. celerio, serpa, para'ena ) A. larva: body scoli not flattened or feather-like; face patterned or plain, but not striped. PUPA: perhaps with some metallic portions, but not totally metallic; head horns, various . . . B B. larva: scoli of A2-8 of similar form, although those of A2, and/or 7 & 8 may be longests. FOODPLANTS: non- rubiaceous C C. larva: pale grey or pale brown with sides of thorax through A2 much darker than rest of body; scolus of A2 long. PUPA: with huge, hook-like dorsal pro- jection from A2; head horns various (A. phylaca aethalia, melanthe, isis, mesentina , delphicola, (l)calliphiclea, (l)abyla) C. larva: dark, or not patterned as above; scolus of A2 short, pupa: hangs tilted with ventral side superior; head horns short points bent sharply to sides ( A . justina, jordani) B. larva: scolus of A2 different in form from scoli of A3-6 (in A . iphicla the difference is merely the swelling at base of A2 scolus) foodplants: rubiaceous D D. larva: scoli of T2 or 3, and A2 & 7 swollen at base; (in A. iphicla, T2 & A2 only); face not patterned; head chalaza-1 not darker than others (don’t known for A. pseudococala). pupa: horns small, triangular “cat- ears” E ♦The reader should bear in mind that in all Adelpha species, the scoli of T1 and A2 are tiny. 1984] Aiello — Genus Adelpha 27 E. larva: scoli of A2 conical, close together, and with fewer spines at base; scoli of A3-6 very short (A. cy there a, salmoneus, pseudo cocala ) E. larva: scolus of A2 not conical, similar to other scoli except for its swollen base; scoli of A3-6 of moderate length; face smoother than other spe- cies due to reduction or loss of most chalazae (A. iphicla ) D. larva: body scoli about same thickness at middle as at base; scolus of A2 strongly arched towards the posterior, thick and densely spined; face patterned; head chalaza-1 dark, and chalazae-3 & 4 pale com- pared to rest of face. PUPA: head horns curved or recurved to sides F F. larva: scoli of T3 and A7 long, slender, and tapered to sharp point; scoli of A7 & 8 differ in form from each other (that of A8 club-shaped). PUPA: head horns sickle-shaped (A. basiloides, phliassa ) F. larva: body scoli not tapered to sharp point; scoli of A7 & 8 similar in form. PUPA: head horns like tiny asymmetrical leaves (A. cocala, leucophthalma ) Group I major characteristics: (1) Larval body scoli flattened; (2) Larva with face striped; (3) Larval head chalaza-1 dark (also true of GROUPS VI & VII); (4) Larvae assume a curled resting position; (5) Pupa shimmering silver; (6) Valves of male genitalia without clunicula; foodplants: Aquifoliaceae, Bombacaceae, Cecropiaceae, Com- bretaceae, Melastomataceae, Rubiaceae, Urticaceae. Two larval types have been observed within this group: A. Final instar larvae with A2 scolus broad, curved posteriorly, and with median spines spreading nr. celerio, serpa. 28 Psyche [Vol. 91 A. Final instar larvae with A2 scolus longer, slender, straight (although directed posteriorly), and with median spines ascend- ing celerio, paraena Pupae vary only slightly, the more apparent differences being in the length, and degree of curvature of the head horns, and in the shape of the dorsal projections on T2 and A2. Head horn shape and size do not appear to correlate with the two larval types seen so far, and perhaps varies even within a species. celerio, Panama (Aiello, Coley) Larvae are dark, mostly black above and green laterally, with pinkish lateral spots on A2-4, 7 & 8; scoli on thorax and A7 & 8 are pale, while that of A2 is black; subspiracular scoli A2-4 are mixed pale lime and bright green. A black subspiracular stripe passes along the thorax and turns sharply upward across A1 and fuses with the dark scolus of A2. A few days before pupation the pinkish lateral marks change to green, as do the scoli of T3 and A7 & 8; the scolus of A2 remains dark. Just prior to pupation larvae fade to a dirty yellow. Larvae rest in the curled position with the posterior end elevated and the white undersurface of A8 & 9 exposed, and in this configuration resemble a bit of fallen moss and lichen mingled. Pupal head horns are long, and slightly curved near their tips. Adults resemble iphicla in wing pattern. nr. celerio, Panama (Aiello) A larva resembling celerio produced, but with the A2 scolus short and broad, a pupa that had short head horns and appeared identical to my nr. paraena. Unfortunately the pupa died and its identity remains a mystery. celerio diademata, Mexico (Comstock & Velazquez, 1960) The illustrations of this larva and its pupa indicate member- ship in GROUP I; the larva is clearly of the celerio type having long narrow scoli, even on A2. The pupa closely resembles that of the celerio reared by me in Panam&; the head horns are long and slightly curved just before their tips. The authors report that their larva faded to yellow just before pupation. 1984] Aiello — Genus Adelpha 29 paraena, Brasil (Moss, 1933) From its illustration, the larva of this species is of the slender scolus type. Moss noted that it was brighter green than serpa and had none of the orange spines. The record of Rubiaceae for paraena is unique for this group. nr. paraena, Panama (Aiello) In pattern, this larva is very similar to A. celerio, but is paler above, being mottled brown and black, with a white dorsal patch joining A6 & 7. The scoli are brown, that of A2 being the darkest; subspiracular scoli A2-4 are pink. The larva turned yellowish just prior to pupation. The pupal head horns are shorter than those of celerio, and curve outward at about their midpoint. Adults resemble celerio so closely that I did not realize they were different species until rearing them, and doubtless they are mingled in many collections. serpa, Brasil (Moss, 1933) The larva of serpa has the scolus of A2 shorter and denser than the other major scoli, and I take it to be of the same general type as celerio. Moss described the scolus of A2 (segment 6 in his system) as being bushy and dark green, and the other major scoli as ochreous (orange). His larva rested with the anterior portion turned, and resembled a bird dropping. serpa, Brasil (Muller, 1886) Muller’s illustrations of serpa show the larval face, and a scolus from A2. The face is clearly striped, and the scolus is short, thick, and has flattened spines overlapping each other; each of these conditions of the scolus is more extreme than in nr. celerio. Group II major characteristics: (1) Larva pale, with sides of thorax through A2 darker than rest of body; (2) Pupa with huge, laterally flattened, hook-like projection from dorsum of A2; (3) Pupal head horns diverse: leaf-shaped, bent triangular, rounded; 30 Psyche [Vol. 91 (4) Valves of male genitalia armed with row of teeth beginning at apex and extending along lower edge; clunicula broad; foodplants: Aquifoliaceae, Bombacaceae, Cecropiaceae, Ulma- ceae, Urticaceae. abyla , Jamaica (Swainson, 1901) There are several things in Swainson’s description that lead me to place abyla tentatively with GROUP II. “On first segments are two brown horns with sharp points bending over the head; from this to the end of the third segment is silvery grey; then two tiny horns bending backward.” “The [pupa] shape is very curious, resembling the pictures of ‘Punch,’ long nose and all.” It appears that larval segments T1 and A1 were overlooked, probably because of their small size and tiny scoli. The silver area would then be bounded by scoli of T2 and A2, suggesting the two-toned larva of GROUP II. The pupal description may be in reference to the long abdominal hook of GROUP II, or it may be an over- reaction to the shorter dorsal projection of a species in some other group (iphicla perhaps). Unfortunately, no foodplant is mentioned. calliphiclea, Paraguay (Jorgenson, 1921) Jorgensen’s description of the pupa as having the abdomen prolonged toward the thorax, forming a large hook, induces me to place calliphiclea in GROUP II: “El torax arriba con una carena alta como el abdomen en el dorso este ultimo hacia el torax prolongado, formando un gran gancho.” In addition, the non-rubiaceous foodplant (Ilex) is consistent with this group. Each of the five species remaining, possess characteristics 1 & 2 above, and there is no doubt in my mind that they are all close relatives. Pupal head horn shape may have taxonomic value within the group but that cannot be determined until a larger number of species has been reared. delphicola , Brasil (Moss, 1933) The pupal head horns curve to the sides and resemble tiny asymmetrical leaves; the hook on dorsal A2 is rounded at the tip. isis, Brasil (Muller, 1886). Muller’s illustration of the pupa is a lateral view so the head horn shape does not show, however, it is stated in the text that 1984] Aiello — Genus Adelpha 31 the horns are small points bent to the sides. The hook on A2 is enlarged slightly at the tip much as in melanthe. melanthe, Panama (Aiello) The larva is grey (darker above) excepting the dorsal thorax through A2, which is dirty yellow, and the sides of that same area, which are dark brown. The scoli are grey with black spine bases and apices. The larva has a frosted appearance partly due to its patterned scoli, but mostly because the entire body is densely clothed with short thick grey setae. It is the only Adelpha species so far that has rounded pupal head horns. The hook on A2 is slightly expanded at the tip. Adults of melanthe look like large salmoneus but the undersurface is quite different. mesentina, Brasil (Moss, 1933) From Moss’s illustrations, mesentina and delphicola larvae are similar, and no doubt these two species are close relatives. As a result of their close resemblance, Moss did not realize that he had two species until the adults emerged. If his illustrations are accurate, then mesentina has a less rounded pupal hook than delphicola, but the head horns are nearly the same. phylaca aethalia, Panama (Aiello, Coley) The larva is greenish or pinkish grey, and speckled with darker grey (especially on the dorsum); the sides of the thorax through A2 are dark brown, and some individuals have a dark brown oval on each side of A5, just below the scolus. Pupal head horns are short triangles bent to the sides. Adults are very close look-alikes of cocala. Group III major characteristics: (1) Larval scoli A2-6 all similar and short; (2) Pupa tilted so that ventral side is superior (at least in justina ); (3) Pupal head horns small triangles bent to sides (also occurs in GROUP II); (4) Valves of male genitalia unarmed (also true of GROUP IV); 32 Psyche [Vol. 91 foodplants: Verbenaceae. The most interesting thing about this group is the fact that the scolus of A2 is just the same as those of A3-6. Both species, thus far included, rest in the front-curved position, as shown in Moss’s illustration of jordani. Also in both, the pupal dorsal A2 projection is a short point, directed anteriorly, and the T2 projection is smaller. This and group IV have similar genitalia, but feed on different foodplant groups. I see them as close but distinct relatives. jordani, Brasil (Moss, 1933) The larva is dull green with black-spined, orange scoli; the pupa is white with black spots. justina, Panama (Aiello & Small) Early final instar larvae are dark brown (lighter dorsally) with paler scoli (black at tips of those on T2 and A7 & 8), but later change to black and white checkered with transverse orange bands joining members of scolus pairs (A2-8). The straw-colored pupa has a dorsal metallic sheen, from head through Al, and lacks the prominent spotting of jordani. Adults of justina closely resemble leucophthalma and the two can be found flying together. Group IV major characteristics: (1) Larva with scoli of A2 conical, close together, and with fewer spines at base; (2) Larval face with chalazae reduced in size; (3) Pupal head horns of this and Group V, are small, triangular “cat-ears;” (4) Valves of male genitalia unarmed (also true of GROUP ill); foodplants: Sabicea (Rubiaceae). nr. cocala, Brasil (Mtiller, 1886) The larva illustrated does not at all resemble that of cocala, so I guess Muller’s label “bei cocala ” to have been based upon adult appearance. Like cytherea, this larva shows the conical scolus of A2, the thick based scoli of T2, and A7, and the very short scoli of A3-6. As well, Muller’s larva fed on Sabicea. 1984] Aiello — Genus Adelpha 33 cytherea, Brasil (Moss, 1933) Of this larva, Moss commented, “Now in two shades of brown, on the stem or near flower-head easily mistaken for a part of the plant. ...” cytherea marcia, Panama (Aiello) The larva is patterned dark and light brown; with dark brown, oblique lateral stripes, which cross segments A3-8. A thin white subspiracular line expands into a lime-green mark on A7-9. The pupa is straw-color but takes on a silver or gold sheen at certain angles. pseudococala, Brasil (Moss, 1933) Moss states merely that the larva of this species is identical to cytherea, although larger and greener at the ends. The pupal abdominal projection is directed towards the posterior as in salmoneus. salmoneus, Panama (Aiello) The dark brown second through fourth instar larvae, with their grey, oblique, lateral stripes, are well camouflaged amongst the dark, pale-veined, young leaves of their foodplant. Fourth and fifth instars have red-brown heads. Early final instar larvae are brownish green, and become greener with age. By late final instar, larvae are bright yellow- green; the dorsal area of A4 & 5 is pale purple-brown; A4 & 5 each bear a dark lateral purple mark; the head is red-brown; scoli of T2 & 3, and A2 & 7 are brown-purple, while that of A8 is still green. The pupa is bronze with dorsal gold, and has an A2 projection that is prolonged more towards the posterior end than the anterior. Group V major characteristics: (1) Larval scoli of T2 and A2 swollen at base, otherwise, all body scoli usually similar in form; (2) Pupal head horns of this and GROUP IV, are small, triangular “cat-ears”; (3) Larval head uniformly dark, and face smooth except for usual pits, and two pairs of reduced chalazae (nos. 2 & 4); 34 Psyche [Vol. 91 (4) Valves of male genitalia armed with apical row of teeth which extend to outside of valve; foodplants: Antirrhoea, Alseis, Bathysa, Calycophyllum, Gon- zalea, Isertia, Rondeletia, Uncar ia (Rubiaceae). iphicla, Panama (Aiello) The color of iphicla larvae varies from dark grey, through golden brown, and red-brown, to almost black, regardless of the foodplant eaten. The head is always smooth (by Adelpha standards) and also varies, from yellow-brown with black punctations to uniform dark brown or black. Final instar larvae show the most pattern (especially light-colored individuals): dark, oblique lateral stripes on A2-5, which terminate part way up the bases of the scoli, and often but not always, white lateral marks on A2, 7 & 8. The pupa varies from waxy white to straw-color or pale brown, and often is partly or entirely burnished gold or silver, especially on dorsal T1 & 3, and Al. Almost always there is a small silver diamond located on the base of each mesothoracic leg. iphicla, Brasil (Moss, 1933) Moss described iphicla larvae as “very similar to cytherea, the two species evidently being closely related, as is also shown by the pupae.” groups IV and V may be related; the “cat-eared” pupae, and oblique-striped larvae of both, seem to hint at that. As well, both groups of larvae display varying degrees of reduction of head chalaze, and swelling of certain scoli bases. However, the genitalia of the two groups are different, and, in iphicla the scoli on A3-6 are longer and the face smoother than in Group IV. And so, for the moment they will be kept separate, but near to each other. Group VI & VII major characteristics: (1) Larva with scolus of A2 arched towards the posterior, thick, and densely spined; (2) Larval face patterned; (3) Larval head chalza-1 dark (also true of GROUP I); 1984] Aiello — Genus Adelpha 35 (4) Valves of male genitalia armed with apical row of 2-6 fairly uniform teeth; (5) Larval head chalazae 3 & 4 pale, and contrasting with face. foodplants: Rubiaceae. These two groups are very closely related and perhaps will be merged when more is known of their larval and pupal diversity. For now they are placed apart on account of the specialized scoli in basiloides, and differences in the form of the pupal head horns. Group VI ADDITIONAL CHARACTERISTICS: (5) Larval scoli of T3 and A7 long, slender, and tapered to sharp point; (6) Larval scolus of A8 club-shaped; (7) Larval face pattern, independent of pits; (8) Pupal head horns sickle-shaped; foodplants: Alibertia, Amaioua, Bertiera, Ixora (all Rubiaceae). basiloides, Panama (Aiello) Larvae are mottled black and brown, and are paler towards the posterior end. The dark and pale portions intersect obliquely along a line beginning at the subspiracular scolus of A4, and terminating dorsally at the beginning of segment A7. The dorsal dark portion thus comes to a point at the beginning of A7. As larvae approach pupation, they may become tinged with rose or green, especially at scoli bases, and laterally on A4-6. The pupa is pearly white with black tiped head horns, and appears empty. Head horns vary somewhat in length and degree of curvature, even among larvae collected on the same plant. Both extremes are illustrated (Figure 6). phliassa, Brasil (Moss, 1933) Comparing my observations of basiloides with Moss’s descrip- tion and illustrations of phliassa, it appears that the two are extremely close relatives. plesaure, Brasil (Muller, 1886) The pupa illustrated appears identical to those of phliassa and basiloides. Hall (1938) feels that plesaure and phliassa are one and the same species. 36 Psyche [Vol. 91 Group VII ADDITIONAL CHARACTERISTICS: (4) Larval face pattern due to dark pits against a paler face; (5) Pupal head horns shaped like tiny asymmetrical leaves (also occurs in GROUP II); foodplants: Calycophyllum, Chomelia, Malanea, Pentagonia, Psychotria, Uncaria, Warscewiczia (all Rubiaceae). cocala, Panama (Aiello) Early fifth instars are crypticaly patterned with golden brown and black; later the pattern remains but the colors become moss- green, black, and cream, and a pinkish grey and black area appears on dorsal A3-6, plus a broad, oblique, lateral pinkish or yellowish stripe across A4 and 5 together. Some individuals have a subspiracular lime-green mark on each of A7 & 8. The pupa is dark green. cocala, Brasil (Moss, 1933) The curved, thick scoli of A2 show clearly in Moss’s illustration, but his larvae had more than one lateral oblique stripe. The pupae appear nearly identical. His record of Emmotum (Icacinaceae) as a foodplant, in addition to Malanea (Rubiaceae) is odd, and is the only reported deviation, by cocala, from Rubiaceae. leucophthalma leucophthalma, Panama (Aiello & Small) The larva of leucophthalma is apparently indistinguishable from that of cocala (G. Small, personal communication) and the two species must be closely related. The pupa also is very similar to cocala, but differs in being brown or copper-color, and in having the abdominal projection rather square, the thoracic projection more pointed, and the head horns farther appart at their bases. leucophthalma leucophthalma (as /. tegeata ), Costa Rica (Young, 1974) The observations by Young are consistent with those of Gordon Small and myself. Species Excluded From This Classification erotia, Brasil (Muller, 1886) An isolated scolus, figured by Muller and labelled erotia, would 1984] Aiello — Genus Adelpha 37 place that species in GROUP I, but I would like to see more evidence. In addition, the erotia wing-undersurface pattern does not have the sharply stamped appearance typical of GROUP I, and it is possible that the butterly was misidentified. iphicla iphicleola, Mexico (Comstock & Vazquez, 1960) This record was in error; the authors misidentified Doxocopa as Adelpha. melona, Brasil (Moss, 1933) The larval description and illustrations in Moss do not give sufficient detail to place this species, although from the pupa it belongs in GROUP VII. thesprotia, Brasil (Moss, 1933) The larval description and illustration in Moss do not give sufficient detail to place this species, although I suspect that it may represent an eighth group. As well, there is a good deal of disagreement concerning the identity of Moss’s thesprotia; Forbes (unpublished) believed that Moss actually had nea, a species which on the basis of genitalia probably belongs to GROUP I. Moss’s statement that the larva rested in a curled position (typical of GROUP I), lends support to Forbe’s suspicion. However, from what can be seen of the scoli in Moss’s illustration, they are all similar in size and form, are not flat, and the pupa looks more like those of GROUP VIII. Significance of Adult Wing Pattern When adult specimens of Adelpha are sorted into the species groups just outlined, the result is a jumble of wing pattern types that would make any biologist wince, this author included. However, it would seem even less natural to split groups that are based upon the presumably more conservative characters of the immatures. I prefer to think that in Adelpha, wing pattern not only does not reflect species relationships, but instead may be intended to deceive. One might expect adult wing characters to be the most specialized and difficult to interpret; adults move about with ease and interact with each other and with other butterfly species, as well as with potential predators, while in most cases the immature stages must contend, in 38 Psyche [Vol. 91 a less active way, with predators and parasites alone. An explana- tion of Adelpha’s wing pattern madness, might be that this genus is comprised of several groups, of various affinities, which together form a large mimicry complex, perhaps based upon members whose larvae feed on alkaloid-bearing plants (e.g., Rubiaceae). Possible examples are found among the unrealted look-alikes reared by me in Panama (Figure 7): phylaca aethalia (non-rubiaceous feeder (NR)) closely resembles cocala (rubiaceous feede (R)); justina (NR) resembles leucophthalma (R); melanthe (NR) is larger than, but very similar in pattern to salmoneous (R); and celerio and nr. paraena (both NR) are very similar in pattern to iphicla and basiloides (both R). The idea is intriguing, but must be labelled “pure speculation” until data on palatability of Adelpha butterflies is available. In its defense, however, I would like to point out that there are several examples of Adelpha look-alikes in uncontestably unrelated butterflies. Nymula velabrum (Riodinidae), whose wing pattern closely resembles that of A. iphicla, and several other Nymula species that are look-alikes of other Adelpha types, all depart from the basic wing pattern of their genus. Female Doxocopa laure and pavon (Nymphalidae) resemble A. iphicla, while female D. Clothilda are even more convincing look-alikes of A. salmoneus. Male Doxocopa Clothilda and pavon differ from the females of their species, and do not resemble Adelpha, while the male of D. laure has an Adelpha- like wing pattern with overlying purple iridescence. Pyrrhogyra hypsenor (Nymphalidae) is similar in appearance to Adelpha species that have broad white bands, and has fooled at least one lepidopterist (Muyshondt, 1974). Old World Relatives The few published accounts of life histories for Old World Limenitidinae provide intriguing glimpses into the trove of informa- tion awaiting the attention of lepidopterists. Many larval and pupal forms, similar to those of Adelpha, occur among Old World genera, and in addition, many utilize the same foodplant genera as Adelpha and display similar larval behavior. Life history studies, that compare Old and New World groups, are essential to our understanding of generic limits and relationships within this subfamily. 1984] Aiello — Genus Adelpha 39 6 Morrell (1954) gives a fascinating account of larval behavior in several Malayan genera of the tribes Limenitidini (Athyma, Moduza, Pandita ), and 7Neptini (Lasippa, Neptis, Phaedyma). Except for Neptis leucoporos, the larvae, of the butterflies he observed, expose the midrib, or a secondary leaf vein, by feeding around it (just as Adelpha does); young larvae rest out on these slender supports. Like Adelpha also, Athyma, Moduza, and Pandita accumulate their feces to form a mass at the base of the support vein; Neptini do not. Judging from Morrell’s illustrations, the larva of Moduza resembles a combination of Adelpha cocala and basiloides, es- pecially in having short thick scoli on A2; its pupa is remarkably like A. cocala or leucophthalma, but with slightly stalked head horns; the host plants are mainly Rubiaceae (Mussaenda, Nauclea, Timonias, Uncaria, Wendlandia), with one record on Oleaceae (Olea). The larvae of Athyma kanwa feeds on Uncaria, and is described by Morrell (1960) as “brown with delicate green branched spines, and resembles a growth of moss on a decaying patch of leaf;” the pupa (illustrated in Morrell, 1954) is silver with long head horns and to me looks for all the world like a pupa of Adelpha celerio. In form, Athyma nefte larvae (Morrell, 1954) are reminiscent of a very plump Adelpha iphicla larva, and have been reported on Glochidion (Euphorbiaceae) and Mussaenda (Rubiaceae); the pupa is “golden brown in colour, with two dorsal plates of brilliant metallic gold,” and from what can be determined from its picture, it looks like the pupa of Adelpha iphicla or cytherea, but with an exagerated A2 projection, and like those species it appears to have short pointed head horns. Pandita sinope is described by Morrell (1954) as having a larva similar to that of Athyma nefte in color and general appearance; the illustration of the pupa is also similar to A. nefte, but has a less 6 According to Corbet and Pendlebury (1978), five of the seven names used by Morrell (1954) should be amended: his Neptis columella = Phaedyma columella; Neptis heliodore = Lasippa tiga; Neptis nata = Neptis leucoporos; Parathyma kanawa = Athyma kanwa; Parathyma nefte = Athyma nefte. His Moduza procris and Pandita sinope remain unchanged. 7Neptini is defined by Eliot (1969) to include Aldania, Lasippa, Neptis, Pantoporia, and Phaedyma. [Vol. 91 Psyche Figure 7 a. Upper and lower surfaces of adult Adelpha species reared in Panam&: BAS = basiloides, CEL = celerio, COC = cocala, CYT = cytherea marcia, IPH = iphicla, JUS = justina, LEU = leucophthalma, MEL = melanthe, nr. PAR = near paraena, PHY = phylaca aethalia, SAL = salmoneus. Species in the upper row are rubiaceous feeders as larvae, whereas larvae of those in the lower row feed on various non-rubiaceous plants. 1984] Aiello — Genus Adelpha 41 Figure 7b. Lower surfaces of adult Adelpha species reared in Panama. Abbreviations as in figure 7a. 42 Psyche [Vol. 91 exagerated A2 projection and thus appears almost identical to pupae of Adelpha ip hie la and cytherea. Larvae of P. sinope feed on Uncaria (Rubiaceae). The valves of the male genitalia in Pandit a sinope and Athyma nefte are unarmed, have cluniculae, and look very similar to those of Adelpha groups III & IV. A number of other life history accounts for Old World Limenitidini have been published, but it is not the intent of this paper to report them in detail. Instead, I merely wish to point out a few of the intriguing similarities that exist between Old and New World groups, in hope of prompting other workers to investigate this promising group. Acknowledgements Many people contributed to this project, and I am indebted to each of them for their enthusiastic help and support. I would especially like to thank Gordon Small for his encouragement during this project, as well as for generous logistical help, identification of butterflies, and for the many valuable larvae which he has brought to me. Richard Vane-Wright of the British Museum provided photographs of a number of Adelpha type specimens, as well as helpful comments about them; Adelpha celerio and phylaca aethalia were identified by comparison with these photographs. Robert Diaz, James Mallet, and Henry Stockwell also located caterpillars for this project, and Bob Diaz and Dagmar Werner contributed their time and effort to translate several German papers. Lissy Coley made specimens, from her own project, available for study. I would like to thank Ricardo Cortez for logistical and technical help, and Robert Robbins and Gordon Small for reading the manuscript. This paper would be deficient without the carefully executed drawings of Marshall Hasbrouck, which were made using a combination of live and preserved material plus photographs, or without the foodplant list compiled by Philip DeVries. Phil has made this list available for use by other lepidopterists, in spite of the fact that it is as yet unpublished. Robert Silberglied gave encourage- ment in the earliest stages of this project. I would like to thank the Image Systems Branch, Tropic Test Center, Corozal, Panama, for the photographs (Figure 7) of adult specimens. Without the facilities and library of the Smithsonian Tropical Research Institute, 1984] Aiello — Genus Adelpha 43 Panama, and the support of National Geographic Society Grant 2444-82, this work would not have been possible. Literature Cited Aiello, A. (Unpub) Insect Rearing Records (mainly for Lepidoptera). Each year is in a separate notebook; lot numbers used in the present paper refer to pages in these notebooks. Barcant, M. 1970. Butterflies of Trinidad and Tobago. 314 pp.; Collins. London. Coley, L. (Unpub) L. Coley has reared a number of Lepidoptera on Ochroma and Cecropia. Numbers given are her identification numbers. Comstock, J. A. and L. VAzquez G. 1960. (1961) Estudios de los ciclos bioldgicos en Lepid6pteros Mexicanos. an inst biol Mexico 31: 349-488. Corbet, A. S., and H. M. Pendlebury 1978. The Butterflies of the Malay Peninsula. 3rd edition revised by J. N. Eliot, xiv + 578 pp., 35 pi., 438 figs. Malayan Nature Society. DeVries, P. J. (Unpub) List of host plants of butterflies of Costa Rica, compiled from his own data plus that of a dozen other workers. The last compilation was prepared during early 1982. Dewitz, H. 1879. Naturgeschichte Cubanischer Schmetterlinge. In: Zeitschrift flir die gesammten Naturwissenschaften. Bd. 52: 166-167. Dornfeld, E. J. 1980. The Butterflies of Oregon, xiv + 276 pp.; Timber Press. Forest Grove, Oregon. Eliot, J. N. 1969. An analysis of the Eurasian and Australian Neptini (Lepidoptera: Nymphalidae). 155 pp., 3 pi., 101 text-figs. Supplement 15 to bull brit mus. Eltringham, H. 1923. Butterfly Lore. 180 pp.; Oxford, at the Claredon Press. Ferris, C. D., and F. Martin Brown (editors) 1980. Butterflies of the Rocky Mountain States, xviii + 442 pp.; University of Oklahoma Press. Norman. Fruhstorfer, H. 1907. Adelpha. pp. 510-533 + pis. 106-1 10a. In: seitz, A. The Macro- lepidoptera of the World. Div. II, Vol. 5, viii + 1139 pp. Godman, F. D., and O. Salvin 1879- 1901. Biologia Centrali-Americana. Insecta. Lepidoptera-Rhopalcera. Vol. I, xiv + 487 pp. (1879-1901); Vol. II, 782 pp. (1887-1901). 44 Psyche [Vol. 91 Hall, A. 1938. On the types of Adelpha (Lep., Nymphalidae) in the Collection of the British Museum, entomologist 71: 184-187, 208-211, 232-235, 257-259, 284-285. Hayward, K. J. 1950. Las especies y formas Argentinas del gfenero Adelpha. acta zool lilloana 9: 375-393. Hoffmann, F. 1937. Beitrage zur Naturgeschichte brasilianischer Schmetterlinge. II. ento- mologisch zeitscrift 51: 212-213, 231-232. Howe, W. H. 1975. The Butterflies of North America, xiii + 633 pp., 97 color plates containing 2093 illus. and 32 text figs. Doubleday & Company, Inc. Garden City, New York. Jones, E. D., and F. Moore 1883. Metamorphoses of Brazilian Lepidoptera from San Paulo, Brazil. Second Series, proc lit philos soc Liverpool 37: 229-259. Jorgensen, P. 1921. Sobre algunos nuevos enemigos de la yerba-mate, Ilex paraguariensis. REV SOC CIENTIFICA PARAGUAY 1(1): 27-30. Kirkbride, A. C. G. 1982. A preliminary phylogeny for the neotropical Rubiaceae. pl syst evol 141: 115-122. Miller, L. D., and J. Y. Miller 1970. Notes on two rare Mexican Adelpha and related Central American species (Nymphalidae). j lep soc 24(4): 292-297. Morrell, R. 1954. Notes on the larval habits of a group of nymphalid butterflies. Malayan nat j 8: 157-164. 1960. Common Malayan Butterflies. Malayan Nature Handbooks, xii + 64 pp., 20 pis. Longman. London. Moss, A. M. 1933. Some generalizations on Adelpha, a neotropical genus of nymphalid butterflies of the group Limenitidi. nov zool 39: 12-20 + pis. 1 & 2. MOller, W. 1886. SUdamerikanische Nymphalidenraupen, versuch eines natlirlichen Sys- tems der Nymphaliden. zool jahrb, zeit syst, geogr biol thiere 1: 417-678 + pl. 12-15. Muyshondt, A. 1974. Notes on the life cycle and natural history of butterflies of El Salvador. V. A. Pyrrhogyra hypsenor (Nymphalidae-Catonephelinae). j ny ent soc 82(3): 163-172. Orsak, L. J. 1977. The Butterflies of Orange County, California, xii + 349 pp.; University of California, Irvine. Center for Pathobiology misc publ No. 3; Museum of Systematic Biology res ser No. 4. 1984] Aiello — Genus Adelpha 45 Peterson, A. 1962. Larvae of Insects: An Introduction to Nearctic Species. Part I. Lepidoptera and Plant Infesting Hymenoptera. Columbus, Ohio. Riley, N. D. 1975. A Field Guide to the Butterflies of the West Indies. 224 pp. + 338 color illus.; Collins. London. Scudder, S. H. 1889. Butterflies their Structure, Changes and Life-histories with special reference to American Forms, viii + 322 pp.; Henry Holt & Co. New York. Silva, A. G. D’A. E., C. R. Goncalves, D. M. Galvao, A. J. L. Goncalves, J. Gomes, M. DO N. Silva, and L. DE. Simoni. 1967- 1968. Quartro Catalogo dos insetos que vivem nas plantas do Brasil, sues parasitos e predadores. parte I (2 vol., 906 pp.) & parte II (2 vol., 887 pp.). Ministerio da Agricultura. Rio de Janeiro, GB, Brasil. Small, G. (Unpub) List of Panamanian Butterflies. This list is based upon twenty years of observations in Panama. Swainson, E. M. 1901. Notes on lepidopterous larvae from Jamaica, B.W.I. j ny ent soc 9: 77-82. Young, A. M. 1974. Notes on the natural history of a rare Adelpha butterfly (Lepidoptera: Nymphalidae) in Costa Rica high country, j ny ent soc 82(4): 235-244. EARWIGS OF THE CALIFORNIA CHANNEL ISLANDS, WITH NOTES ON OTHER SPECIES IN CALIFORNIA (DERMAPTERA)* By Scott E. Miller Museum of Comparative Zoology Harvard University Cambridge, Mass. 02138 Although the earwigs of the California Channel Islands were included in Langston and Powell (1975) and Langston and Miller (1977), newly accumulated records extend their distributions signifi- cantly (Table 1). Some of these new records are due to increased collecting activity by interested entomologists, but most probably represent range expansion by the earwigs, aided by increased human activity on the islands. The two species involved, Euborellia annu- lipes (Lucas) (Carcinophoridae) and Forficula auricularia Linnaeus (Forficulidae), are both considered to be introduced to California (Langston and Powell 1977). The two are readily distinguished by the shape of the forceps (Langston and Powell 1977: figs. 3, 11), as well as the adults being wingless (E. annulipes ) or winged (F. auricu- laria). This note is based on the collections of the California Academy of Sciences (CAS), Natural History Museum of Los Angeles County (LACM), Peabody Museum of Yale University, San Diego Natural History Museum (SDNHM), Santa Barbara Museum of Natural History (SBMNH), Smithsonian Institution (USNM), and University of California at Berkeley (UCB), as well as my own fieldwork on all islands from 1976 to 1984. Euborellia annulipes (Lucas) This cosmopolitan species has been established in coastal south- ern California since at least the 1880s (Langston and Powell 1977). Channel Islands records are as follows: Santa Rosa: A single female was taken in July 1939 (LACM). A colony was found at Johnsons Lee in May 1977 (SBMNH). Santa Catalina: A series was collected at Avalon in October 1908 (USNM), but recently only single indi- * Manuscript received by the editor June 26, 1984. 47 48 Psyche [Vol. 91 viduals have been collected at Catalina Harbor (July 1979, LACM) and on the beach at Toyon Bay (May 1981, SBMNH). San Cle- mente: A series was collected in 1885 or 1886 by T.L. Casey (USNM), one was collected in November 1941 (LACM), and it has been taken in the 1970s around Wilson Cove (military living com- pound) and the new airport. Forficula auricularia Linnaeus This cosmopolitan species has not been established in California as long as E. annulipes and has undergone rapid expansion of range in the state in the last fifty years (Langston and Miller 1977). It was probably not established in coastal southern California until at least the 1940s (Langston and Powell 1975). Channel Islands records are as follows: Santa Rosa: Colonies sampled during the 1970s at Beechers Bay (ranch headquarters), Arlington Canyon, and Wreck Canyon (CAS, SBMNH, USNM). Santa Cruz: Powell (1981) dis- cusses the establishment of this species between the late 1960s and 1978. It is now well established in the Central Valley. Santa Cata- lina: Between 1975 and 1983, it has been collected at Catalina Har- bor, Cherry Cove, Cottonwood Canyon, Gallaghers Beach, Parsons Landing, Twin Harbors (Cat Harbor), and Toyon Bay (LACM, SBMNH), and can be very abundant (S.G. Bennett pers. comm.). San Nicolas: Taken in 1980 and 1982 at the military living com- pound (LACM, SBMNH). San Clemente: First taken at the new airport in March 1972 (UCB), it was taken at Wilson Cove and Lemon Tank in December 1981 (SBMNH). Santa Barbara: One female taken in June 1983 (LACM). Discussion Both species are generally restricted to the areas around human activity (i.e. ranch and military facilities) and beaches. The earwigs are omnivorous, sometimes predaceous on other insects, often feed- ing on live or dead vegetation, often injuring crops on the mainland (Langston and Powell 1975). These species, especially Forficula auricularia, can also be household nuisance pests, and can cause considerable annoyance in large numbers. Based on the data presented above, E. annulipes has been present on three islands for 40 to 100 years and maintains small popula- 1984] Miller — Earwigs of Channel Islands 49 Table 1. Summary of earwig distribution on the California Channel Islands. Island Euborellia annulipes Forficula auricularia San Miguel Santa Rosa X X Santa Cruz X Anacapa Santa Catalina X X San Nicolas X San Clemente X X Santa Barbara X tions, probably near the sites of initial introduction by man. F. auricularia, however, has apparently become established on six islands, mostly during the 1970s. It has not yet been recorded from San Miguel and Anacapa islands, but should be expected there. The agencies which administer the islands should attempt to discourage the rapid expansion of F. auricularia, by preventing additional introductions and eradicating existing populations when possible. Appendix The following are significant additions to Langston and Powell (1975) and Langston and Miller (1977): Following Brindle (1971) and Steinmann (1975), the proper name of Spongophora apiceden- tata Caudell is Vostox apicedentatus (Caudell). Euborellia femoralis (Dohrn), a new California record, was reported from Red Hill Ma- rina, Calipatria, Imperial County, by Steinmann (1981). Forficula auricularia has been established in the San Diego area, new south- ern record in California, since at least the mid 1970s (specimens in SDNHM). Acknowledgements I thank the National Park Service, U.S. Navy, Santa Cruz Island Co., Vail and Vickers Co., and Santa Catalina Island Conservancy for permission to collect on the islands; those and the Catalina Island Marine Institute, Santa Barbara Museum of Natural His- tory, Southern California Academy of Sciences, TRW, and the 50 Psyche [Vol. 91 University of California at Santa Barbara Marine Science Institute for logistic and financial support of surveys; the curators of the collections consulted; and S.G. Bennett, P.W. Collins, F.G. Hoch- berg, C.L. Hogue, R.W. Klink, R.L. Langston, C.D. Nagano, J.A. Powell, H. Steinmann, and D.B. Weissman for specimens and assistance. Literature Cited Brindle, A. 1971. A revision of the Labiidae (Dermaptera) of the Neo-tropical and Nearc- tic Regions. III. Spongiphoridae. J. Nat. Hist. 5: 521-568. Langston, R.L. and S.E. Miller 1977. Expanded distribution of earwigs in California (Dermaptera). Pan-Pac. Ent. 53: 114-117. Langston, R.L. and J.A. Powell 1975. The earwigs of California (Order Dermaptera). Bull. Calif. Ins. Surv. 20: 1-25. Powell, J.A. 1981. Five insects believed to be newly established or recolonized on Santa Cruz Island, California (Dermaptera, Lepidoptera). Bull. Soc. Calif. Acad. Sci. 79: 97-108,(1980). Steinmann, H. 1975. A Survey of the Neotropical Vostox Burr species (Dermaptera: Labii- dae). Acta Zool. Acad. Sci. Hung. 21: 435-445. 1981. A study of the circumtropical Dermaptera material in the “Instituut voor Taxonomische Zoologie”, Amsterdam. Acta Zool. Acad. Sci. Hung. 27: 187-210. NESTING BIOLOGY AND DEFENSIVE BEHAVIOR OF MISCHOCYTTARUS (MONOCYTTARUS) MEXICANUS CUBICOLA (VESPIDAE:POLISTINAE)* By Henry R. Hermann and Jung-Tai Chao Department of Entomology, University of Georgia, Athens, Georgia 30602 Introduction Mischocyttarus mexicanus (de Saussure) is one of two species in this genus that occurs in the U.S. (Krombein et al., 1979). This genus is primarily Neotropical, being treated taxonomicaily by Bequaert (1933), Richards (1945, 1978) and Zik&n (1935, 1949). Litte (1977, 1979) described the biology of M. mexicanus in Florida and M, flavitarsis (de Saussure) in Arizona. M. m. mexicanus is found in Texas, Mexico and other parts of Central America. M. m. cubicola to date has been found in Georgia, Florida, Alabama, Cuba and the Bahamas (Krispyn and Hermann, 1977; Krombein et al, 1979). Litte’s study of the nesting biology and behavior of M. m. cubicola (Litte, 1977, then described as M. mexicanus) was carried out at Archbold Biological Station at Lake Placid in southern Florida (Highlands County) where weather conditions allowed the wasps to nest throughout the winter. Our primary study area in Georgia had a more temperate climate where a 3 to 4 month hibernation period was necessary. Sapelo Island, a 14-mile-long barrier island in McIntosh County, Georgia, consists of Holocene and Pleistocene deposits (Duncan, 1982). There is no official weather station on Sapelo Island. However, the average annual rainfall from 1944 to 1964 at McKinnon Airport on the southern end of St. Simons Island, about 7 miles south of Sapelo, was 134.6 cm, with half of this falling during a 4-month period from June to September. Mean minimum temperature for the coldest months was about 6.1° C (43° F). Temperatures as low as 0° C (32° F) occur only 15 days of the year. The lowest recorded temperatures were —10° C 0 Manuscript received by the editor February 6, 1984. 51 52 Psyche [Vol. 91 (14° F) in December, 1962, and —7.8° C (18° F) in February, 1958. Mean daily maximum temperature during the summer was under 32.2° C (90° F), only 49 days per year having a maximum of 32.2° C or above. The highest recorded temperature was 38.9° C (102° F) in June, 1950. The purpose of this paper is to report the general nesting biology and defensive behaviors of M. m. cubicola at Sapelo Island. Materials and Methods Four trips were made to Sapelo Island (fall of 1982; late winter- early spring, 1983; June 9-12, 1983, August 23-30, 1983), one to Patterson Island, Ga. (adjacent to Sapelo) (June 11, 1983), one to Orlando, Polk County, Fla. (Aug. 7, 1983) and one to Haines City, Polk County, Fla. (Aug. 5-8, 1983). Two of the trips to Sapelo Island were preliminary surveys of the island while the later trips to Sapelo and the other three study areas revealed nests in various stages of development. When an active nest was found, an attempt was made to provoke adults to defend by prodding them with the tip of a pen or comparable instrument. Defensive behaviors were video taped (Hitachi VT-650A VTR and VK-C600 color camera) during August so that they could be examined closely and repeatedly. Due to their readiness to escape under provocation in the early nesting season (June), some adults were lost. Following a study of their defensive behaviors, the nest and remaining adults were taken. Nest contents and information on attending adults were later recorded. A comparison between paper surfaces of young and older nests was made through the use of scanning electron microscopy. Parts of nests were removed and adhered to specimen stubs with double-stick tape. They were coated with palladium-gold for 2 minutes in a Hummer sputter coater and observed with a Cambridge Mark II scanning electron microscope. Results and Discussion Nesting Sites Nests of M. m. cubicola were found on buildings, other man- made structures and on vegetation. Nests on the former were most often found on the wooden frames of windows, on the wooden support beams of metal roofed outbuildings and on the underside of 1984] Hermann & Chao — Mischocyttarus 53 Figures 1 and 2. Nests of M. m. cubicola at Sapelo Island, showing the general structure and safety area on the back of the nest (arrow). concrete arbors. Small (5-15 cells) uninhabited nests (Figs. 1 and 2) were numerous on the wooden frames of windows of the R. J. Reynolds home adjacent to the Marine Institute dormitory at the southern end of the island. Uninhabited nests were also found less abundantly on wooden structures of other buildings at the island’s south end. Nest abandonment at these locations was partially due to occasional sprayings of pesticides. Three nests were found in metal and ceramic bell-like wind chimes in the Haines City, Polk County, Fla., location. Nests on vegetation on Sapelo Island were found only on the underside of the leaves (Fig. 3) of ihe cabbage palm, Sabal palmetto (Walt.). This tree and Serenoa repens (Bartr.) (the common saw palmetto) (Fig. 4) are the most predominant members of the palm family, Arecaceae (= Palmae), found on Sapelo Island. One nest of this wasp was also collected from S. palmetto on Patterson Island, but nests were not found on buildings in that location. No nests were found on saw palmetto, Serenoa repens , on either island. In contrast to our findings, Litte (1977) found nests of this wasp almost exclusively on the leaves of Serenoa repens. Nests were found on cabbage palm on an island in a man-made lake near Orlando, Fla., and a single nest was found on the leaf of a plant in the genus Heliconia (Musaceae) at that location. Bequaert (1933) mentioned a nest found in Cuba that was built on Spanish moss ( Tillandsia , Bromeliaceae) about six feet above the ground at the periphery of a swampy margin of a stream. 54 Psyche [Vol. 91 Sabal palmetto (Fig. 3), most abundant on the south end of Sapelo Island, appears ideal for a nesting site because of the way in which mature leaves hang in a horizontal plane, each leaf offering a pair of nesting sites, one on each side of the midrib. Serenoa repens, on the other hand, has leaves that extend upward from the ground in a more-or-less vertical plane (Fig. 4). Such leaves offer very little protection from both weather and birds. Reproductive Potential Based on information from 10 nests, a single egg layer was apparently on each nest during the June 9-12 observation period (Table 1). This is based entirely on ovary size. Although escaping females were lost, the most gravid females of other polistine species (Hermann, unpublished) usually remain on the nest when provoked while less gravid females depart. Therefore, it is our opinion that each nest had a single egg layer. Only in one case (Nest #4) a second female on the nest had other than small, non-reproductive ovaries, and they were developed but not large. At least half of the nests found at Sapelo Island, and probably more, were in the predaughter (= preemergence) phase. Litte (1977) also found her nests at this stage with a single egg layer. Females other than the gravid queen on nest #4 in which there were no pupae are considered to be cofoundresses. This supports the pleometrotic status of this subspecies as found earlier by Litte (1977). Our data (Table I) do no show a significant difference in wing length between the queens and non-queens (X2 0.5 (1) = 3.841). Predation and Potential Predators A high degree of predation by birds of wasp nest immatures was apparent on Sapelo Island. Although bird species were not identified in this location, nests were often found shredded and abandoned, a condition characteristic of bird damage. Numbers of damaged nests were not recorded. Jeanne (1975), Richards and Richards (1951), and Turillazzi (1984) have stated that birds and ants are the most important predators of social wasps in general. Birds apparently are the most significant predators in temperate zones (Jeanne, 1975). Jeanne (1970) reported predation on Mischo - cyttarus by bats. Litte (1977) reported seeing blue jays attacking nests of M. m. cubicola in south Florida and she listed the Carolina wren, scrub jays, common yellowthroat, cardinal, thrashers, mockingbird and 1984] Hermann & Chao — Mischocyttarus 55 Figures 3 and 4. Plants found to be nesting sites for M. m. cubicola by us and by Litte (1977). 3, Sabal palmetto, showing draping nature of the leaves, making it an ideal site for nesting wasps. 4, Serenoa repens, showing the nature of growth so that leaves do not drape, making them a poor site for nests. robin as potential vertebrate predators. She also pointed out that scrub jays and red headed woodpeckers are known predators of Polistes adults. Invertebrate predators of wasp immatures were Dorymyrme x flavopectus, Crematogaster ashmeadi, Pheidole flori- dana, Camponotus floridanus, and Monomorium floricola (Formi- cidae). Spiders were found to catch and feed on wasp adults. Starr (1981) reported blue jays and summer tanagers as possible polistine predators. Litte (1977) found that defense against vertebrate attack was ineffective, while defense against ants was effective in most cases. Polistine wasps are apparently free of most terrestrial predators while some vespines are heavily preyed upon by them (Preiss, 1967). Details of Nest Structure and Architectural Defense Of the ten nests that were collected between June 9 and June 12, 1983 (Table I), six had 15 cells or more. The average number of cells was 13.90 (SD — 8.41). Although there is no information about colony growth rate at this time, it appears that most nest initiation occurred in May. Data collected from nests 1, 3, 5 and 7 (Table I) with less than 10 cells imply that either: 1) they were reinitiated after predation or 2) they were late initiators. Further study on these possibilities may give us a better understanding of the selection forces acting upon developing colonies. Most nests in June collections had a single female that dominated the face of the nest while other adult nest occupants spent much of their time on the Table I. Nest structure, nest inhabitants and reproductive condition of females of Mischocyttarus m. cubicola at Sapelo Island, June 9-12, 1983. Total Wing Nest # of # of adult length # Cells (1) females Eggs Larvae Pupae Female Ovaries (mm) 56 Psyche [Vol. 91 in m on a\ o ggo £ ^ oo O ON o o J Q NO (N o o . „ J J i—I J J O 01 < < < n 3 3 •r > OO On oo I-- m oo NO 3 3 o (U 3 « M 3 oj 3 "C m O £ Z i © § § S u 8 £? . o "3 O 8.® ® v 01 ^ > ©« 01 T3 «n — ON 13 o i ii O (4i £ 7? S -5 S .s .22 oo oo 3 00 „o oi > . CO a a § 01 o 1 o C/2 Q _ 1 ° X* 3 iS C .5 TJ is s ci 1/3 8 2 52 c8 g 3 ° 3 « § f c/3 C I £ o 3 00 "7 3 3 •a © 0 c « a £ ■o u 5 >1 '% © c/3 ® ii T3 1 8. I 2 Pu « -• oi S 73 8 2 & i T3 (« 00 2 00 03 01 g 2 5 3 PU B 6 .8 .§ X) E 8.5 2 2 3 _ £ .5 O' w o 13 00 ^ 3 .5 (4) 3 00 3 « C o £ © > « 13 x » - o o <2 1 ^ £ o' O ON 1984] Hermann & Chao — Mischocyttarus 57 back of the nest. There is a significant positive correlation (r = 0.095, p <0.001) between the number of females and the number of cells in a nest (Table I). Small nests that were relatively new were light gray in color, while larger nests were often a darker, brownish color. The walls of small, young nests were translucent and the outline of a larva could be seen inside. In addition, the paper was loosely made (Fig. 8) so that occasional holes existed in the cell walls. The loose appearance and light color of the paper indicated that very little if any head secretions were added other than what was necessary to adhere the layers together. Also, very little mandibular working of the paper was done by the wasps at this time. Both paper darkening and smoothing (Figs. 6 and 7) appear to commence between the 4-5 cell stage and the time of capping. Cell cap darkening following spinning is done by adults. Freshly spun caps were whitish while those of older pupae had brownish colored wood particles adhering to them (Fig. 5). Darkening of the caps and the addition of wood particles blended the caps with the rest of the nest, making them cryptic. This is a primary defense (Edmunds, 1974). Another primary defense is seen in the way in which most of the nests hang on their pedicels. Some are elongate, pendulous (Fig. 1), others more round in appearance (Fig. 2). Irrespective of shape, all were constructed with the nest back away from the direction of provocation. This is most apparent on nests constructed on buildings. This allows adults to retreat from provocation rather than defend. Nests on Sahal palmetto often were horizontally elongate, following the available space on the underside of the leaf. Most successful nests were constructed under leaves that had additional folds which allowed further protection from birds. Nests found in bell-like wind chimes in Haines City, Florida, were exceptionally well protected. The bells, in effect, were analogous to the bag-like (calyptodomous) nests of arboreal vespines. Predation on these nests by birds would be difficult. Similar and other forms of architectural defense have been reported from other social hymenopterous species by numerous individuals and has been reviewed by Hermann and Blum (1981). 58 Psyche [Vol. 91 Figures 5-8. Scanning electron micrographs of paper from nests of M. m. cubicola. 5, Cap on cell of a late pupa (nest #9, June collection, 31 cells), showing wood and other debris deposited by adult wasps, thus giving the cap a cryptic appearance (50X). 6, Paper on outside of same nest, showing its non-woody, well- worked surface (500X). 7, Pedicel of same nest, showing extremely smooth texture (100X). 8, Pedicel of smaller, newer nest (nest#l, June collection, 44 cells), showing loose nature of woody constituents (100X). Defensive Behavior other than Architectural Young Colonies Attack behavior as studied by numerous individuals and reviewed by Hermann and Blum (1981) appeared to be non-existent in this species during early colony life. However, a pseudoattack was elicited from wasps upon strong provocation. Strong provocation was carried out by tapping on the leaf upon which the nest was built or by actually touching the nest. Pseudoattack is defined as adult flight from the nest in the direction of the intruder. This differs from escape in which flight is not directed toward the intruder. Pseudoattack is readily demonstrated in other local polistines, e.g., 1984] Hermann & Chao — Mischocyttarus 59 Polistes annularis (Linnaeus) and P. exclamans (unpublished). We would classify pseudoattack as a very strong warning behavior. Pseudoattack was best demonstrated by M. m. cubicola in more mature colonies (found in August on Sapelo Island), while escape was more prevalent in younger colonies. Other warning behaviors are as follows, some of which have been described by Jeanne (1972), Litte (1977, 1979), Starr (1981) and Windsor (1972) for this and other species of Mischocyttarus. Defensive Posture — antennae forward, wings raised, body lifted and mandibles sometimes spread. Strong provocation was necessary to elicit this combination of behaviors in young nests. Litte (1979) reported that M. flavitarsis males and females both assume a defensive posture when disturbed. Increased Movement on the Nest — increased movement gives the nest an agitated appearance. Increased provocation elicits increased movement by adult wasps and apparently lowers the threshhold for attack. Wing Raising — generally with approximately an angle of 57° - 107° between wings (X = 84.6, SD = 13.7), and a mean angle of 38° above the horizontal plane of the body. While wing raising is one of the first warning behaviors elicited in most polistines (Hermann and Blum, 1981; Starr, 1981; West Eberhard, 1969), most females of M. m. cubicola on young nests did not raise their wings unless they were strongly provoked. Wing raising also has been reported for M. drewseni by Jeanne (1972) and for M. m. cubicola by Starr (1981). Raised wings appear to have two separate positions, a smaller angle of separation (approximately 60°) for wasps that are weakly provoked and a greater angle of separation (approximately 90- 100°) for wasps that are strongly provoked. However, there is considerable overlap in these two positions. Wing Buzzing — a prolonged wing movement, not generally demonstrated in early colonies except under strong provocation. Wing buzzing was demonstrated by M. flavitarsis (Litte, 1979), but adults did not attack in the pre-emergence phase. Attack was prevalent following wing buzzing in the post-emergence phase, however. Wing fluttering — sporadic rapid flapping of wings, of a much shorter duration than buzzing. Strong provocation was necessary to elicit this behavior in young nests. Wing fluttering frequencies may be found in Table III. 60 Psyche [Vol. 91 Forward Jerking and Mandibular Pecking — a rapid movement forward and a biting at the object of intrusion. Pecking was also demonstrated in the absence of forward jerking. Although pecking behavior was easily elicited in a young colony, it appeared to be demonstrated in such cases almost entirely by the queen. Pecking is a behavior commonly used by adult females toward small hymenop- terous parasitoid intruders. In P. annularis, forward jerking and pecking have been observed by us to chase and/or discard intruding ants from the nest. We witnessed such behavior in older colonies observed in August, 1983. Backward Jerking — best demonstrated under strong provocation. Abdominal Pumping — although abdominal pumping, a rhythy- mic extension and retraction of the gaster, is seen as normal behavior in all vespids, it appears to be exaggerated during provocation. Abdominal twisting — this behavior was not evident on a young nest except under very strong provocation. However, when strongly provoked, most females of a young nest demonstrated this behavior as well as exaggerated abdominal pumping. Movement to back of nest (retreat) — a behavior that was demonstrated upon continuous provocation. Even females that demonstrate warning behaviors move to the back of the nest upon continuous provocation. Escape — this is readily demonstrated in young colonies under provocation. In fact, it is difficult to elicit defensive behaviors in young colonies and collect all of the adults because of their readiness to escape. Nest departure by adults of more mature colonies is expressed more as pseudoattack in which case adults return to the nest after a short erratic flight. Adults in escape flights do not return for a longer period of time. Leg waving — this behavior was demonstrated on occasion in young colonies and readily in mature ones. Although females of young colonies are reluctant to leg wave unless strongly provoked, this behavior was less difficult to elicit in a young colony than were some of the other behaviors. Leg waving frequencies may be found in Table III. None of the behaviors appeared to be demonstrated in a particular sequence, and many of the behaviors were demonstrated by some individuals and not by others. Escape and retreat were the most dependable behaviors in young (preemergence) colonies. 1984] Hermann & Chao — Mischocyttarus 61 Escape and pseudoattack also occur in colonies of P. annularis, P. exclamans and P. fuscatus during the preemergence period (un- published). Mature Colonies A mature colony is defined here as one in which daughters have emerged (post-emergence colony), and defensive behaviors are readily expressed, many of them requiring little provocation. Although there is very little difference in nest size between nest 9 of Table I (June collection) and nest 1 of Table II (August collection), defensive behavior in August was considerably more evident. The mean number of cells of the 4 nests collected in August (Table II) was 72.5 (SD = 46.3). Few males were found to have emerged (nests 2, 4, Table II). By comparing the colonies in June and August, we noticed that not only the colony structure changes significantly but the defensive behaviors demonstrated by colony members are also quite different. The change in defensive behaviors could be: 1) a seasonal, physiological change in adults, or 2) related to investment, such as higher total number of immatures, more older larvae and pupae, and more nesting material. There was an apparent synchrony of movement among warning females toward a moving intruder, due in part to a periodic hooking together of their tarsi and legs. This behavior is difficult to describe and would have gone unnoticed had it not been recorded on video tape and played back at a slower speed. A sequence of wing raising, leg waving and wing fluttering was readily apparent. Under strong provocation, wing fluttering, leg waving and exaggerated abdominal pumping were demonstrated simultaneously. The frequency of movement in leg waving was Table II. Nest structure and inhabitants of nests of Mischocyttarus m. cubicola at Sapelo Island, Georgia, and Haines City, Florida, August 5-8, 23-30, 1983. Nest # # of Cells Adults Males Females Eggs Larvae Pupae 1(HC) 40 0 14 8 22 10 2(HC) 42 1 15 8 13 14 3(HC) 56 0 7 23 21 1 4(S) 152 3 27 16 60 38 HC, Haines City, Florida; S, Sapelo Island, Georgia. Mean number of cells = 72.5 (SD = 46.3). 62 Psyche [Vol. 91 Table III. Statistical Analysis of Frequencies of Readily Expressed Warning Behaviors in Colonies of M. m. cubicola (1) Exp. # Leg Waving Movements/ Sec. Wing Fluttering Beats/ Sec. Pseudoattack Flight Time in Sec. 1 8.3 5.6 2.7 2 9.1 6.3 3.3 3 7.7 6.7 1.6 4 9.1 6.3 1.6 5 9.1 5.3 5.8 6 7.1 7.7 3.7 7 7.7 5.3 2.8 8 9.1 5.6 3.7 9 7.1 6.3 9.0 10 7.7 6.7 1.6 X, SD, CV X = 8.2 SD = .801 CV = 0.64 X = 6.18 SD = .72 CV = .51 X = 4.18 SD = 2.5 CV = 0.60 (1) Behaviors were recorded from different females. 7. 1-9.1 /second (X = 8.2, SD = .801, CV = .64)_ (Table III). The frequency of wing fluttering was 5. 3-7. 7/ second (X = 6.2, SD = .72, CV = .51). We found no correlation (r = .07) between movement frequencies of leg waving and wing fluttering in different females (Table III). It appeared to us that a threshold had to be reached before leg waving or wing fluttering would come about, and once this threshold was reached, the duration of each behavior was dependent upon the degree of continued stimulus. However, very little variation appeared to exist in the frequencies of these behaviors in relation to the excitedness of the colony (Table III). Pseudoattack by multiple females was easily elicited. Yet, attempts at stinging were relatively rare. Departing females flew in the direction of intrusion but most often culminated their pseudo- attack with an erratic, vertically undulating flight in their return to the nest. In recording the duration of 10 randomly selected pseudoattack flights, each flight usually lasted only 1. 6-9.0 seconds (X = 4.2, SD = 2.5, CV = 0.60) (Table III). Behavior after returning to the nest often involved rapid walking on the nest face in apparent examination of the nest surface and its coinhabitants. The checking of cells and cell contents was not as 1984] Hermann & Chao — Mischocyttarus 63 apparent as has been reported by Starr (1981) for some other polistine species. At times, females would spin in place on the nest immediately upon returning from a pseudoattack flight. Spinning is described as a rapid turning of the body throughout a small radius of movement. Also, post-attack grooming is sometimes evident. Nests 1-3 from Haines City, Florida, reacted to provocation much like nest 4 (Table II), even through there was a considerable difference in nest size. One apparent behavioral difference existed. Nests from Haines City were adjacent to one another in metal and ceramic bell-like wind chimes. When strongly provoked, adults that left the nest in pseudoattack at times returned to a neighboring nest rather than to their own and rapidly moved around the nest face. Observing this indicates to us that possibly the three nests were initiated by siblings. We have observed similar behavior in Polistes annularis. Comparisons between June and August nests point out that defensive behavioral changes occur in M. m. cubicola as a function of time. This is also true for P. annularis, P.fuscatus and P. exclamans (unpublished) and appears to be widely recognized for other vespid species (Hermann and Blum, 1981). The major differences occur in the defensive attitude of eusocial wasps between the pre- and post-emergence periods. Summary Mischocyttarus mexicanus cubicola on Sapelo Island, Georgia, nests on buildings, other man-made structures and on the underside of leaves of the Cabbage Palm, Sabal palmetto. Although defensive behaviors expressed in young colonies at first appear to be poorly demonstrated in this subspecies, they include nest architecture and, under strong provocation, pseudoattack and subsequent erratic flight, general nest excitability, defensive posture, wing raising, wing buzzing, wing fluttering, forward jerking, mandibular pecking, backward jerking, abdominal pumping, abdominal twisting, retreat and escape. None of the warning behaviors were demonstrated in young colonies in a consistent manner or in a particular sequence. The most consistent behaviors in young colonies were retreat and escape, whereas in mature colonies all of the warning behaviors recorded for young colonies were expressed readily and in a predictable sequence. 64 Psyche [Vol. 91 Acknowledgements We would like to extend our appreciation to the following individuals for their warm hospitality during this investigation: Mr. and Mrs. Ralph Talarico, Haines City, Florida; Mr. and Mrs. Fred Woodward, Patterson Island, Georgia. Also, we would like to thank the University of Georgia Marine Institute at Sapelo Island for their support in this project. References Bequaert, J. 1933. The Nearctic social wasps of the subfamily Polybiinae (Hymenoptera: Vespidae). Entomol. Amer. 13:87-148. Duncan, W. H. 1982. Vegetation of Sapelo Island. Georgia Dept. Nat. Res. Publ. 75 pp. Edmunds, M. 1974. Defence in Animals. Harlow, Essex: Longman, 357 pp. Hermann, H. R. and M. S. Blum 1981. Defensive mechanisms in the social Hymenoptera. [In] Social Insects, H. R. Hermann, ed., Academic Press, New York, vol. 2, pp. 77-197. Jeanne, R. L. 1970. Chemical defense of brood by a social wasp. Science 168:1465-1466. 1972. Social biology of the Neotropical wasp Mischocyttarus drewsenii. Bull. Mus. Comp. Zool., Harvard Univ. 144:63-150. 1975. The adaptiveness of social wasp nest architecture. Q. Rev. Biol. 50:267-287. Krispyn, J. W. and H. R. Hermann. 1977. The social wasps of Georgia: Hornets, yellowjackets, and polistine paper wasps. USDA Agric. Res. Bull. 207, 39 pp. Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and B. D. Bucks. 1979. Catalog of Hymenoptera in America North of Mexico. Smithsonian Inst. Press, Washington, D. C. vol. 2, p. 1516. Litte, M. 1977. Behavioral ecology of the social wasp Mischocyttarus mexicanus. Behav. Ecol. Sociobiol. 2:229-246. 1979. Mischocyttarus flavitarsis in Arizona: Social and nesting biology of a polistine wasp. Zts. Tierpsychol. 50:282-312. Preiss, F. J. 1967. Nest site selection, microenvironment and predation of yellowjacket wasps, Vespula maculifrons (Buysson) (Hymenoptera, Vespidae) in a decidous Delaware woodlot. M. Sc. thesis, Univ. Delaware, 81 pp. Richards, O. W. 1945. A revision of the genus Mischocyttarus de Saussure (Hymen., Vespidae). Trans. R. ent. Soc. Lond. 95:295-462. 1978. The Social Wasps of the Americas. Fletcher and Son Ltd, Norwich. 1984] Hermann & Chao — Mischocyttarus 65 Richards, O. W. and M. J. Richards. 1951. Observations on the social wasps of South America (Hymenoptera: Vespidae). Trans. R. ent. Soc. Lond. 102:1-170. Snelling, R. M. 1981. Systematics of social Hymenoptera. [In] Social Insects, H. R. Hermann, ed., Academic Press, New York, vol. 2, pp 369-453. Starr, C. K. 1981. Defensive tactics of social wasps. Doc. Dis., Univ. Georgia, Athens, 108 pp. Turillazzi, S. 1984. Defensive mechanisms in Polistes wasps. [In] Defensive Mechanisms in Social Insects, H. R. Hermann ed., Praeger Scientific, New York (in press). West Eberhard, M. J. 1969. The social biology of polistine wasps. Univ. Mich. Mus. Zool. Misc. Publ. 140:1-101. Windsor, D. M. 1972. Nesting association between two Neotropical Polybiine wasps (Hymen- optera, Vespidae). Biotropica 4:1-3. ZikAn, J. F. 1935. Die sozialen Wespen de Gettung Mischocyttarus Saussure, nebst Beschreibung von 27 neuen Arten (Hym., Vespidae) Archos. Inst. Biol. Sao Paulo 1:143-203. 1949. O genero Mischocyttarus Saussure (Hymenoptera, Vespidae), con a descripao de 82 species novas. VBoln. Parq. nac. Itatiaia 1:1-251. REVISION OF THE MILVINA GROUP OF THE WOLF SPIDER GENUS PARDOSA (ARANEAE: LYCOSIDAE)i By C. D. Dondale and J. H. Redner Biosystematics Research Institute Research Branch, Agriculture Canada, Ottawa, Ontario, Canada K1A OC6 Introduction The genus Pardosa C. L. Koch is a large and widespread group of wolf spiders. Several species groups have been recognized in North America (Lowrie and Dondale 1981). The objective of the present paper is to define and revise the milvina group. The milvina group of the genus Pardosa consists of 18 American species of wolf spiders. The greatest concentration of species seems to be in the southeastern United States and Mexico. Only P. milvina (Hentz), P. saxatilis (Hentz), and P. littoralis Banks range northward as far as Canada, and only P. portoricensis Banks, P. hamifera F. Pickard-Cambridge, and P. littoralis have been found in the West Indies. P. fastosa (Keyserling) extends into South America as far as Ecuador. None are found on the U.S. Pacific coast or in the U.S. Rocky Mountains, and few specimens are recorded from the Great Plains. Terminology for anatomical parts is defined by Dondale and Redner (1978) and by Figures 2-7 and 41-45 here. Measurements are given as the mean and standard deviation for 10 to 20 specimens or as the range for fewer. Relationships The external genitalia, in our opinion, provide the best indicators of relationships within the milvina group. The inferred relationships among the species are shown in Figure 1. Females of all species in the group have the lateral margins of the median septum raised and thickened at the site of the copulatory openings (character 1, Fig. 1; Fig. 45). This modification appears to enlarge the openings and to reinforce the margins, and we speculate 1 Manuscript received by the editor December 10, 1983. 67 DIT 68 Psyche [Vol. 91 Cladogram of the milvina group. See text for definition of the numbered components. 1984] Dondale & Redner — Genus Pardosa 69 that each opening may accommodate not only the embolus, which is a slender shaft, but the enlarged, sclerotized tip of the conductor as well. The condition is not found elsewhere in Pardosa except in two species of the Old World wagleri group, which have a quite different kind of median septum, hood, and copulatory tubes. We infer the condition found in the milvina group to be apomorphic, and the absence of the condition to be plesiomorphic. The base of the median septum is more or less rectangular in females of certain species of the group (character 4, Fig. 1; Fig. 50). This condition is unique among the species of Pardosa, and is inferred to be apomorphic. The plesiomorphic state is a triangular base. Also, a strongly tapered anterior part of the septum (character 9, Fig. 1; Fig. 65), in combination with a single hood cavity, is inferred to be apomorphic and a broader anterior part to be plesiomorphic. The epigynal hood may extend posteriad at the sides, thus defining a raised, tapered, median area (character 12, Fig. 1; Fig. 58). This condition, found only in specimens of the saxatilis complex, is inferred to be apomorphic. The simpler, lobed hood found in the other members of the milvina group is inferred to be plesiomorphic. The median apophysis, which is believed to assist indirectly in aligning the embolus tip with the copulatory opening, may be unusually short and blunt (character 2, Fig. 1; Fig. 3), with its basal process located at midlength on the apophysis (character 5, Fig. 1; Fig. 8) and having its mesal margin thickened (character 7, Fig. 1; Fig. 8). These states are inferred to be apomorphic and the more slender, elongate apophysis, with the basal process located at the base and with the mesal margin not thickened, to be plesiomorphic. The conductor in the milvina group is a stout rod, lying transversely in the groove between the apical division of the genital bulb and the tegulum. In the unexpanded bulb, only its tip is visible (Fig. 3), but it becomes fully exposed by dissection (Fig. 6). The apomorphic states of this character are inferred to be “angulate or pointed on basal margin” (character 3, Fig. 1; Fig. 6), “fluted at tip” (character 6, Fig. 1; Fig. 11), “possession of dark, shiny knob near tip” (character 8, Fig. 1; Fig. 21), and “knob excavated” (character 10, Fig. 1; Figs. 38, 40, arrows). The plesiomorphic states are inferred to be “straight basal margin”, “thick tip”, “knob lacking”, and “knob not excavated”. 70 Psyche [Vol. 91 Certain segments of the male palpus may be covered dorsally with reflective white setae (characters 11, 11a, Fig. 1; Figs. 16- 18). These setae probably serve as a visual cue to the female during male courtship, and their presence is believed to be apomorphic. In saxatilis, only the patella bears such a covering; in atlantica, both patella and tibia are covered, and in parvula, both segments plus the basal half of the cymbium are covered. Males of only the remotely related P. tesquorum (Odenwall) have a similar covering on the palpal patella. We infer that the presence of this covering in the milvina group is apomorphic, and that the plesiomorphic state of the character is the involvement of a single segment. Unidentified Species A name associated in the past with the milvina group, namely, Lycosa canadensis Blackwall, 1871, treated as a junior synonym of P. milvina by Chamberlin (1908), could not be identified by us, and is treated here as nomen dubium. The holotype of cana- densis was an unidentifiable juvenile and is no longer in exist- ence. Pardosa accurata Becker, 1886, described from “Mexico”, has been identified as an unknown species of Lycosa, s.l, through examination of the holotype by us; the type is a juvenile speci- men deposited in the Institut Royal des Sciences Naturelles de Belgique. Description Total length 3.00 to 7.50 mm. Carapace smoothly convex at sides, vertical anteriorly at sides, approximately uniform in height between dorsal groove and posterior row of eyes, covered sparsely with short setae; eye area black, often iridescent; anterior row of eyes somewhat procurved, distinctly shorter than middle row, with median eyes equal in size to lateral eyes or somewhat larger and lo- cated slightly closer to laterals than to each other. Legs moderately long and strong, with thin scopulae and sparse setae; femur I with 3 dorsal macrosetae, 2 prolaterals (near tip), 2 or 3 retrolaterals; tibia 1 with 2 slender bristlelike dorsals, 2 prolaterals, 2 retrolaterals, 3 pairs of ventrals; basitarsus I with 1 bristlelike dorsal, 3 prolaterals, 2 or 3 retrolaterals, 3 paired and 1 unpaired ventral; trochanters with deep notch distally at tip. Abdomen ovoid, covered with dense, short setae and with cluster of longer, erect setae at anterior end. 1984] Dondale & Redner — Genus Pardosa 71 Male palpus usually black and contrasting with pale areas on legs and carapace, often with fringes of black setae at sides of segments; palea (Fig. 6) projecting liplike at tip of genital bulb, giving rise prolaterally to slender, rather short, slightly curved embolus and giving rise basally to elongate, stout conductor that lies largely hidden between base of palea and tegulum; terminal apophysis toothlike or bladelike, located retrolaterodistally (Fig. 6); median apophysis short, straight or somewhat curved, with strong basal process (Fig. 3). Epigynum (Fig. 41) rather long (often longer than twice maximum width of median septum), with median septum broad posteriorly and slender or evanescent anteriorly; hood small, distinct, with single cavity; copulatory openings with margins raised and thickened. Copulatory tubes (Fig. 44) short, angulate or somewhat curved, with swelling on ventral or lateral side; sperma- thecae small, bulbous. KEY TO MALES OF THE MILVINA GROUP 1. Distal process of median apophysis short, broad, lacking hook at tip (Fig. 3) 2 I'. Distal process of median apophysis elongate, usually hooked at tip (as in Fig. 9, arrow) 4 2(1). Terminal apophysis broad (ventral view, Figs. 2, arrow, 5); conductor smoothly curved on basal margin (Fig. 5) bellona Banks 2'. Terminal apophysis more slender (ventral view, Figs. 6, 7); conductor angular or pointed on basal margin (Figs. 6, 7, arrows) 3 3(2'). Conductor with hooked point on basal margin (Fig. 6); embolus bent near tip (Fig. 6) delicatula Gertsch and Wallace 3'. Conductor with angulate basal margin (Fig. 7); embolus gently curved at tip (Fig. 7) hamifera F. Pickard-Cambridge 4(1'). Median apophysis with basal process small, located about midlength of apophysis (Fig. 8); conductor thin, fluted at tip, lacking knoblike process at tip (Fig. 11) 5 4'. Median apophysis with basal process located at base of apophysis or, if located at middle then large (Figs. 19, 24); 72 Psyche [Vol. 91 conductor thickened at tip, bearing dark, shiny, knoblike process near tip (as in Figs. 21, 26, 38) 8 5(4). Median apophysis with distal process straight at tip (Fig. 8, arrow) sagei Gertsch and Wallace 5'. Median apophysis with distal process hooked at tip (as in Figs. 9, 10, 14) 6 6(5'). Terminal apophysis separated from mesal swelling by acute angle (Fig. 12); median apophysis with distal process strongly hooked (Fig. 9, arrow) fastosa (Keyserling) 6'. Terminal apophysis separated from mesal swelling by broad curve (Fig. 13, arrow); median apophysis with distal process weakly hooked (Fig. 10) 7 7(6'). Median apophysis with marginal swelling broad (Fig. 10, arrow) desolatula Gertsch and Davis 7'. Median apophysis with marginal swelling narrow (Fig. 14, arrow) mayana sp.n. 8(4'). Patella (and sometimes additional segments) of palpus covered dorsally with reflective white setae (most notice- able in living or dried specimens) (Figs. 16-18); conductor sinuous along basal margin (Fig. 21, arrows) 9 8'. Patella of palpus covered dorsally with dark setae (though few reflective white setae may be present); conductor with single curve along basal margin (as in Figs. 25, 38) . . . 11 9(8). Tibia (as well as patella, and sometimes basal half of cymbium) covered dorsally with reflective white setae (Figs. 17, 18) 10 9'. Tibia of palpus covered with dark setae (Fig. 16) .... saxatilis (Hentz) 10(9). Basal half of cymbium (except narrow band at basal margin) covered dorsally with dark setae (Fig. 17).... atlantica Emerton 10'. Basal half of cymbium covered with reflective white setae (Fig. 18) parvula Banks 11(8'). Terminal apophysis large, extending basad to or beyond tip of embolus (Figs. 22, arrow, 25) littoralis Banks 11'. Terminal apophysis smaller, extending only short dis- tance basad (as in Figs. 24, 35, 42) 12 1984] Dondale & Redner — Genus Pardosa 73 12(1 T). Terminal apophysis arched retrolaterally (Figs. 23, arrow, 26); median apophysis small, occupying about one-third length of genital bulb (Fig. 23) saltonia sp.n. 12'. Terminal apophysis arched mesally, or not arched (Figs. 27, 34, 40); median apophysis larger, occupying distinctly more than one-third of genital bulb (Figs. 24, 31, 42). 13 13(12'). Terminal apophysis with mesal swelling toothlike (Fig. 27, arrow); median apophysis thick throughout most of its length, with basal process swollen (Fig. 24) pauxilla Montgomery 13'. Terminal apophysis with mesal swelling not toothlike, i.e., lacking sharp point (Fig. 34); median apophysis slender at middle (Figs. 28, 37) 14 14(13'). Median apophysis with distal process expanded (ventral view, Fig. 28, arrow). Dorsum of abdomen with pale median band flanked by paired distinct, dark, longi- tudinal bands portoricensis Banks 14'. Median apophysis with distal process more slender (Figs. 31, 37, 42). Dorsum of abdomen dark or mottled, or, if pale mesally then lacking paired dark longitudinal bands 15 15(14'). Terminal apophysis broad, blunt (Fig. 30, arrow). Species occurring only north of Tropic of Cancer (Map 5) . . . milvina (Hentz) 15'. Terminal apophysis more slender, pointed (Figs. 33, 36, 39). Species occurring only south of Tropic of Cancer (Map 6) 16 16(15'). Median apophysis abruptly angled on mesal margin (Fig. 35, arrow). Carapace width less than 1.35 mm. Dorsum of abdomen with median band of reflective white setae . . guadalajarana sp.n. 16'. Median apophysis curved on mesal margin (Figs. 37, 42). Carapace width greater than 1.35 mm. Dorsum of abdomen lacking median band of reflective white setae 17 17(16'). Terminal apophysis rather long (Fig. 36). Tibia and basitarsus I with fringe of long, erect setae along prolateral and retrolateral surfaces. Carapace with black lateral margins longivulva F. Pickard-Cambridge 74 Psyche [Vol. 91 17'. Terminal apophysis shorter (Fig. 39). Tibia and basitarsus I lacking fringe. Carapace usually with pale lateral margins marialuisae sp.n. KEY TO FEMALES OF THE MILVINA GROUP OCCURRING IN EASTERN CANADA AND UNITED STATES 1 . Median septum extending anteriad nearly to level of hood (Figs. 46, 60, 65) 2 F. Median septum extending anteriad one-half length of epigynum or less (Figs. 58, 70) 4 2(1). Median septum broad anteriorly (Fig. 60, arrow) littoralis Banks 2'. Median septum distinctly tapered anteriorly (Figs. 46, 65) 3 3(2'). Median septum with expanded posterior part concave at lateral margins (Fig. 65, arrow) pauxilla Montgomery 3'. Median septum with expanded posterior part convex at lateral margins (Fig. 46, upper arrow) delicatula Gertsch and Wallace 4(1'). Hood continuing posteriad at sides where it defines a raised, tapered median area (Fig. 58, arrows) 5 4'. Hood continuing posteriad at sides where it defines a depressed, non-tapered area (Fig. 70, arrow) milvina (Hentz) 5(4). Species restricted to Great Lakes-St. Lawrence region and southward in Appalachian Mountains (Map 3) saxatilis (Hentz) 5'. Species restricted to southeastern coastal plain (including Florida) and the Mississippi basin (Map 3) 6 6(5'). Species restricted to Florida and eastern Gulf Coast region (Map 3) parvula Banks 6'. Species restricted to Atlantic coast States east of the Appalachians and in the Mississippi basin (Map 3) . . . atlantica Emerton KEY TO FEMALES OF THE MILVINA GROUP OCCURRING IN SOUTHWESTERN UNITED STATES, MEXICO, AND CENTRAL AMERICA 1. Median septum with expanded posterior part approxi- mately rectangular (Figs. 50, 52, 54, 56) 1984] Dondale & Redner — Genus Pardosa 75 desolatula Gertsch and Davis, mayana sp.n., sagei Gertsch and Wallace, and fastosa (Keyserling) 1'. Median septum with expanded posterior part more or less triangular (Figs. 48, 62, 68) 2 2(F). Median septum broad anteriorly, bordered by pair of ridges that extend posteriad along interior surface of septum (Figs. 41, arrow, 43, 44) bellona Banks 2'. Median septum slender or evanescent anteriorly, lacking bordering ridges 3 3(2'). Epigynum with curved ridges posteriorly (Figs. 46, lower arrow, 48, 72, 74, 76) 4 3'. Epigynum lacking curved ridges (Fig. 62) saltonia sp.n. 4(3). Curved ridges concealing lateral angles of median septum (Fig. 46, lower arrow) delicatula Gertsch and Wallace 4'. Curved ridges not concealing lateral angles of median septum 5 5(4'). Epigynum short (ratio of epigynal length to greatest median septum width less than 2:1) hamifera F. Pickard-Cambridge 5'. Epigynum longer (ratio of epigynal length to greatest median septum width more than 2:1) 6 6(5'). Carapace width usually less than 1.6 mm. Retromargin of chelicera with 2 teeth guadalajarana sp.n. 6'. Carapace width usually greater than 1.6 mm. Retro- margin of chelicera with 3 teeth 7 7(6'). Copulatory tubes with small swelling at base (Fig. 75). Carapace margins dark longivulva F. Pickard-Cambridge 7'. Copulatory tubes with large swelling on lateral margins (Fig. 77, arrow). Carapace margins usually pale marialuisae sp.n. KEY TO FEMALES OF THE MILVINA GROUP OCCURRING IN THE WEST INDIES 1. Median septum rather broad anteriorly (Fig. 60, arrow) littoralis Banks 1'. Median septum distinctly tapered anteriorly (Figs. 48, 68) 2 76 Psyche [Vol. 91 2(1'). Copulatory tubes broad, and spermathecae widely sepa- rated (Fig. 49). Epigynum long (ratio of epigynal length to greatest median septum width more than 2:1) hamifera F. Pickard-Cambridge 2'. Copulatory tubes slender, and spermathecae narrowly separated (Fig. 69, arrows). Epigynum shorter (ratio of epigynal length to greatest median septum width less than 2:1) portoricensis Banks Pardosa bellona Banks Figures 2, 5, 41, 43, 44; Map 1 Pardosa bellona Banks, 1898:275, Fig. 21 (pi. 16). Two male and two female syntypes from Coral de Piedras, Baja California (Eisen and Vaslit), deposited in MCZ, examined. Syntypes from San Miguel de Horcasitas, Magdalena Island, and from San Jose del Cabo, deposited in California Academy of Sciences, presumed destroyed. Gertsch 1934:21. Gertsch and Wallace 1935:3, Fig. 14. Roewer 1954:189. Bonnet 1958:3359. Male. Total length 4.50 ±0.34 mm; carapace 2.28 ±0.18 mm long and 1.75 ±0.13 mm wide (20 specimens). Carapace with dark orange median and submarginal areas, and with pair of broad, dark longitudinal bands bordering pale median area; margins dark. Sternum yellow orange, sometimes with several small black spots. Chelicerae orange brown to black, pale mesally; retromargin with 3 teeth. Legs yellow orange; femur I often black basally; tibia and basitarsus I with sparse fringe of long lateral setae. Abdomen pale mesally, dark laterally; venter orange yellow. Terminal apophysis broad; median apophysis short, broad, with large basal process, lacking hook at tip of distal process (Fig. 2); conductor rounded on basal margin (Fig. 5). Female. Total length 4.85 ±0.35 mm; carapace 2.34 ±0.17 mm long and 1.86 ±0.13 mm wide (18 specimens). General color and structure as in male but dark pigment on carapace margin may break into spots, femur I lacking dark pigment at base, and leg I lacking fringe of long lateral setae. Median septum rather broad posteriorly, little tapered anteriorly, flanked anteriorly by ridges that continue along internal surface (Fig. 41); copulatory tubes rather thick, with swelling on ventrolateral surface (Figs. 43, 44). Diagnosis. Specimens of P. bellona most resemble those of P. delicatula and P. hamifera, differing by the broad terminal 1984] Dondale & Redner — Genus Pardosa 77 apophysis, by the rounded basal margin of the conductor, and by the anteriorly broad median septum with paired ridges. Range. Utah south to Colima, Mexico. Included, but not mapped here, are the following: Utah: Monroe Canyon, 112° 10'W, 38°38'N. Nevada: Las Vegas. California: Seeley, 7 mi. west of El Centro; Indian Wells, Riverside Co. Natural History. Males have been collected in December to February, and June to September, females in January, March, June to September, and November. Females with egg sacs were collected in January, March, and May to July. Pardosa delicatula Gertsch and Wallace Figures 3, 6, 45-47; Map 1 Pardosa pauxilla: Montgomery 1904:268 (part, not lectotype). Pardosa delicatula Gertsch and Wallace, 1935:4, Figs. 13, 17. Male holotype and female allotype from Edinburg, Hidalgo Co., Texas (Stanley Mulaik), deposited in AMNH, examined. Gertsch and Davis, 1940: 5. Roewer 1954:189. Bonnet 1958:3365. Male. Total length 4.78 ±0.32 mm; carapace 2.59 ±0.15 mm long and 1.90 ±0.14 mm wide (20 specimens). Carapace with dark orange median and submarginal areas, and with pair of dark brown longitudinal bands flanking pale median area; margins dark, pale, or marked with series of 3 or 4 dark spots. Sternum orange yellow, sometimes suffused with black or with faint, V-shaped dark mark. Chelicerae variable in color; retromargin with 3 teeth. Legs yellow orange; femur I dark prolaterally at base. Abdomen dull yellow mesally, darker laterally, sometimes black throughout; venter pale. Terminal apophysis rather slender (ventral view); median apophysis short, broad, with large basal process, lacking hook at tip of distal process (Fig. 3); conductor with sharp, hooked point on ventral surface; embolus bent near tip (Fig. 6). Female. Total length 5.73 ±0.79; carapace 2.63 ±0.20 mm long and 2.02 ±0.23 mm wide (20 specimens). General color and structure as in male but carapace paler, usually having pale margins. Median septum strongly tapered anteriorly, lacking lateral ridges, convex along lateral margins of posterior part, partly concealed laterally by curved ridges (Fig. 46); copulatory tubes rather slender (Fig. 47). 78 Psyche [Vol. 91 Map 1. Collection localities for Pardosa bellona (squares), P. delicatula (stars), P. hamifera (half-circles), P. littoralis (circles). Diagnosis. Specimens of P. delicatula most resemble those of P. bellona and P. hamifera but differ by the sharp, hooked point on the conductor, by the bent embolus, and by the concealment of the lateral angles of the median septum by curved ridges. Range. Oklahoma and Mississippi south to Chihuahua and Nuevo Leon, Mexico. Natural History. Males and females have been collected in every month except January, and females with egg sacs were taken from April to October. Common habitats are pastures and other grasslands, where they are taken in numbers by pitfall trap; one large collection was taken by sweep net in Louisiana sweet potato fields at night. 1984] Dondale & Redner — Genus Pardosa 79 Pardosa hamifera F. Pickard-Cambridge Figures 4, 7, 48, 49; Map 1 Pardosa hamifera F. Pickard-Cambridge, 1902:320, Fig. 4, 4a (pi. 31). Holotype male from Guatemala (Sarg), deposited in BM(NH), examined. Roewer 1954:187. Bonnet 1958:3373. Pardosa delicata Gertsch, 1934:20. Holotype female from La Zacualpa, Chiapas, Mexico, August 1909 (A. Petrunkevitch), deposited in AMNH, examined. Allotype male and paratype female from the type locality, with same data as for holotype, not examined. Gertsch and Wallace 1935:3, Fig. 18. Roewer 1954:186. Bonnet 1958:3365. NEW SYNONYM. Male. Total length 4.54 ±0.38 mm; carapace 2.36 ±0.18 mm long and 1.83 ±0. 14 mm wide (20 specimens). Carapace with dark orange brown median and submarginal areas, and with dark longitudinal bands flanking median area; lateral margins usually pale. Sternum dark orange, sometimes paler or darker mesally. Chelicerae variable in color; retromargin with 3 teeth. Legs dark orange; femur I often dark at base. Abdomen black with dull yellow heart mark, or yellow brown mesally and darker laterally; venter pale, sometimes suffused with black. Terminal apophysis slender (ventral view); median apophysis short, broad, with large basal process, lacking hook at tip of distal process (Fig. 4); conductor angled on basal margin (Fig. 7). Female. Total length 5.10 ±0.48 mm; carapace 2.45 ±0.22 mm long and 1.95 ±0.19 mm wide (20 specimens). General color and structure as in male but paler. Median septum slender anteriorly, lacking lateral ridges, concave at lateral margins of expanded posterior part (Fig. 48); copulatory tubes rather thick throughout, with ventral swelling (Fig. 49). Diagnosis. Specimens of P. hamifera most resemble those of P. bellona and P. delicatula but differ in having an angular basal margin on the conductor, a gently curved embolus (as in bellona but not as in delicatula ), and in having a slender anterior part and fully exposed posterior part of the median septum. Range. Nuevo Leon south to Honduras; Jamaica, Haiti. Natural History. Males have been collected in Feburary, March, June, and August to December, females in all months except April and September. Habitat is unrecorded. 80 Psyche [Vol. 91 Pardosa sagei Gertsch and Wallace Figures 8, 11, 50, 51; Map 2 Pardosa sagei Gertsch and Wallace, 1937:1, Figs. 1,2. Holotype male and allotype female from El Volcan, Chiriqui, Panama, 26 February 1936 (W. J. Gertsch), deposited in AMNH, examined. Three paratype males and four paratype females from the type locality, with same data as holotype, deposited in MCZ, examined. Roewer 1954:188. Bonnet 1958:3420. Male. Total length 5.00 ±0.35 mm; carapace 2.69 ±0.12 mm long and 2.40 ±0.10 mm wide (20 specimens). Carapace yellow orange mesally and submarginally, with pair of dark longitudinal bands flanking mesal area; margins dark. Sternum orange yellow, sometimes with few black spots. Chelicerae yellow orange; retro- margin with 3 teeth. Legs yellow orange; tibia and basitarsus I with fringe of long, erect, lateral setae. Abdomen yellow brown mesally, darker laterally; venter pale yellow, sometimes with few small black spots. Terminal apophysis small, pointed (ventral view); median apophysis with distal process elongate and straight, with basal process small, located about midlength on apophysis (Fig. 8); conductor thin, fluted at tip, lacking knoblike process (Fig. 11). Female. Total length 5.38 ±0.57 mm; carapace 2.63 ±0.18 mm long and 2.04 ±0.18 mm wide (20 specimens). General color and structure as in male but leg I lacking fringe of setae. Median septum with posterior part approximately rectangular (Fig. 50). Copulatory tubes rather thick, curved near middle (Fig. 51). Diagnosis. Specimens of P. sagei most resemble those of P. fastosa, P. desolatula, and P. mayana. Males of sagei are diagnosed by the straight distal process of the median apophysis. Diagnostic characters are not available for female sagei; the known females were collected together with males. Range. Panama. Natural History. Males and females have been collected in February, March, July, and August. Egg sacs were found in July and August. Pardosa fastosa (Keyserling) Figures 9, 12, 52, 53; Map 2 Lycosa fastosa Keyserling, 1877:618, Figs. 5, 6 (pi. 1). Six syntype males and four syntype females from “Umgebung von St. Fe de Bogota”, Bogota, Colombia 1984] Dondale & Redner — Genus Pardosa 81 (Lindig), deposited in BM(NH), examined. Pardosa fastosa: Petrunkevitch 1911:570. Roewer 1954:186. Pardosa fastuosa: Bonnet 1958:3367 (incorrect subsequent spelling). Pardosa uncatula F. Pickard-Cambridge, 1902:319, Figs. 27, 28 (pi. 30). Lectotype male from La Palma, Costa Rica (Tristan), deposited in BM(NH), examined and here designated. Three paralectotype males and six paralectotype females from the type locality, with same data as lectotype, deposited in BM(NH) and here designated. One male and two females from original syntype series, collected at the type locality and with same data as lectotype, described herein as P. mayana new species. Banks 1909:219. Gertsch 1934:20 (part; proposed synonymy of uncatula with pauxilla, later disclaimed by Gertsch and Wallace (1935)). Roewer 1954:190 (part). Bonnet 1958:3427 (part). NEW SYNONYM. Male. Total length 5.26 ±0.39 mm; carapace 2.73 ±0.22 mm long and 2.09 ±0.17 mm wide (20 specimens). Carapace with orange yellow median and submarginal areas, and with paired dark longitudinal bands flanking median area; lateral margins dark. Sternum black, with pale mesal band anteriorly. Chelicerae dark orange, with black lines; retromargin with 3 teeth. Legs yellow orange; femora I and II dark at base; tibia and tarsus I with lateral fringe of long setae. Abdomen black, with dull yellow heart mark; venter yellow, reticulated with black. Terminal apophysis stout, toothlike, connected to mesal swelling by acute angle (ventral view); median apophysis with distal process strongly hooked at tip (Fig. 9); conductor thin, fluted at tip, lacking knoblike process at tip (Fig. 12). Female. Total length 5.57 ±0.54 mm; carapace 2.70 ±0.23 mm long and 2.11 ±0.20 mm wide (20 specimens). General color and structure essentially as in male, but leg I lacking fringe and black pigment, and pale median area of abdominal dorsum extended posteriad as series of chevrons. Median septum with posterior part approximately rectangular (Fig. 52); copulatory tubes stout, diverg- ing (Fig. 53). Diagnosis. Males of P. fastosa most resemble those of P. sagei, P. desolatula, and P. mayana, but differ in having the terminal apophysis connected to the swelling lying mesal to that apophysis by an acute angle rather than by a curve, and by the strongly hooked tip of the median apophysis. Females of fastosa are anatomically indistinguishable from those of sagei, desolatula, and mayana, but most specimens examined were accompanied by males, or origi- nated in localities from which no males of the other species were known. 82 Psyche [Vol. 91 Map 2. Collection localities for Pardosa sagei Map 3. Collection localities for Pardosa saxatilis (circle), P. (square), P. fastosa (circles), P. desolatula (half-circles), atlantica (stars), P. parvula (half-circles). Question marks indi- P. mayana (stars). cate localities for females of uncertain identity; Nebraska locality was confirmed by discovery of males after the paper was submit- ted for publication. 1984] Dondale & Redner — Genus Pardos a 83 Range. Costa Rica to Ecuador. The record of uncatula from Panama by Petrunkevitch (1925) was not confirmed. The type of Lycosa fastosa viota Strand, 1914, was not found. Natural History. Males have been collected in every month except February and May, and females in every month except May. Females with egg sacs were collected in March, April, July, August, and October to December. Specimens have been recorded from 1,000 to 3,200 meters elevation. Pardosa desolatula Gertsch and Davis Figures 10, 13, 54, 55; Map 2 Pardosa desolatula Gertsch and Davis, 1940:5, Fig. 22. Holotype male and 1 paratype female from Ciudad Victoria, Tamaulipas, Mexico, 12 June 1936 (L. I. Davis), deposited in AMNH, examined. Allotype female from Tamazunchale, San Luis Potosi, Mexico, 25 November 1938 (A. M. and L. I. Davis), deposited in AMNH, examined. Roewer 1954:186. Vogel 1967:104. Male. Total length 3.82-4.55 mm; carapace 2.05-2.42 mm long and 1.53-1.87 mm wide (5 specimens). Carapace with yellow orange to orange brown median and submarginal areas, and with paired dark longitudinal bands flanking pale median area; lateral margins pale to dark. Sternum yellow orange. Chelicerae orange brown to brown; retromargin with 3 teeth. Legs orange yellow; femur I dark on basal half to two-thirds. Abdomen dull yellow brown mesally, darker laterally; venter pale. Terminal apophysis connected to mesal swelling by broad curve; median apophysis with marginal swelling broad, and with distal process weakly hooked (Fig. 10); conductor thin, fluted at tip, lacking knoblike process at tip (Fig. 13). Female. Total length 4.50 ±0.60 mm; carapace 2.17 ±0.19 mm long and 1.65 ±0.17 mm wide (10 specimens). General color and structure essentially as in male, but femur I not black. Median septum with posterior part essentially rectangular (Fig. 54); copulatory tubes stout, diverging (Fig. 55). Diagnosis. Males of P. desolatulata most resemble those of P. sagei, P. fastosa, and P. mayana, but can be distinguished by the broad marginal swelling and weak hook on the median apophysis and by the broad curve by which the terminal apophysis is connected to the mesal swelling. Females are not distinguishable 84 Psyche [Vol. 91 anatomatically from those of sagei, fastosa, or mayana, but all of those examined were accompanied by males, or originated in localities from which no males of the other species were known. Range. San Luis Potosi and Tamaulipas, Mexico. Natural History. Males have been collected in March, June, and November, and females in March, April, September, and Novem- ber. Pardos a mayana sp.n. Figures 14, 15, 56, 57; Map 2 Pardosa uncatula F. Pickard-Cambridge, 1902:319 (part, not lectotype nor para- lectotypes). Type Material. Holotype male from 2 miles north of Fortin de las Flores, 97°01'W, 13°56'N, Veracruz, Mexico, 5 August 1966 (Jean and Wilton Ivie), deposited in AMNH. Three paratypes from La Palma, Costa Rica, deposited in BM(NH). Two paratypes from Honduras and Guatemala, deposited in MCZ. Eighty-seven para- types from Mexico, Guatemala, and Honduras, deposited in AMNH. Male. Total length 4.47 ±0.26 mm; carapace 2.40 ±0.11 mm long and 1.84 ±0.08 mm wide (20 specimens). Carapace with yellow orange median and submarginal areas, and with paired dark longitudinal bands flanking median area; lateral margins pale or dark. Sternum yellow orange. Chelicerae yellow brown; retro- margin with 3 teeth. Legs yellow orange; femur I sometimes black basally; tibiae and tarsi I with fringe of long lateral setae. Abdomen dull yellow brown mesally, darker laterally; venter pale. Terminal apophysis connected to mesal swelling by broad curve; median apophysis with narrow marginal swelling and with distal process weakly hooked (Fig. 14); conductor thin, fluted at tip, lacking knoblike process at tip (Fig. 15). Pardosa desolatula Gertsch and Davis Figures 10, 13, 54, 55; Map 2 Pardosa desolatula Gertsch and Davis, 1940:5, Fig. 22. Holotype male and 1 paratype female from Ciudad Victoria, Tamaulipas, Mexico, 12 June 1936 (L. I. Davis), deposited in AMNH, examined. Allotype female from Tamazunchale, 1984] Dondale & Redner — Genus Pardosa 85 San Luis Potosi, Mexico, 25 November 1938 (A. M. and L. I. Davis), deposited in AMNH, examined. Roewer 1954:186. Vogel 1967:104. Male. Total length 3.82-4.55 mm; carapace 2.05-2.42 mm long and 1.53-1.87 mm wide (5 specimens). Carapace with yellow orange to orange brown median and submarginal areas, and with paired dark longitudinal bands flanking pale median area; lateral margins pale to dark. Sternum yellow orange. Chelicerae orange brown to brown; retromargin with 3 teeth. Legs orange yellow; femur I dark on basal half to two-thirds. Abdomen dull yellow brown mesally, darker laterally; venter pale. Terminal apophysis connected to mesal swelling by broad curve; median apophysis with marginal swelling broad, and with distal process weakly hooked (Fig. 10); conductor thin, fluted at tip, lacking knoblike process at tip (Fig. 13). Female. Total length 4.50 ±0.60 mm; carapace 2.17 ±0.19 mm long and 1.65 ±0.17 mm wide (10 specimens). General color and structure essentially as in male, but femur I not black. Median septum with posterior part essentially rectangular (Fig. 54); copulatory tubes stout, diverging (Fig. 55). Diagnosis. Males of P. desolatulata most resemble those of P. sagei, P. fastosa, and P. mayana, but can be distinguished by the broad marginal swelling and weak hook on the median apophysis and by the broad curve by which the terminal apophysis is connected to the mesal swelling. Females are not distinguishable anatomatically from those of sagei, fastosa, or mayana, but all of those examined were accompanied by males, or originated in localities from which no males of the other species were known. Range. San Luis Potosi and Tamaulipas, Mexico. Natural History. Males have been collected in March, June, and November, and females in March, April, September, and Novem- ber. Pardosa mayana sp.n. Figures 14, 15, 56, 57; Map 2 Pardosa uncatula F. Pickard-Cambridge, 1902:319 (part, not lectotype nor para- lectotypes). Type Material. Holotype male from 2 miles north of Fortin de las Flores, 97°0TW, 13°56'N, Veracruz, Mexico, 5 August 1966 (Jean 86 Psyche [Vol. 91 and Wilton Ivie), deposited in AMNH. Three paratypes from La Palma, Costa Rica, deposited in BM(NH). Two paratypes from Honduras and Guatemala, deposited in MCZ. Eighty-seven para- types from Mexico, Guatemala, and Honduras, deposited in AMNH. Male. Total length 4.47 ±0.26 mm; carapace 2.40 ±0.1 1 mm long and 1.84 ±0.08 mm wide (20 specimens). Carapace with yellow orange median and submarginal areas, and with paired dark longitudinal bands flanking median area; lateral margins pale or dark. Sternum yellow orange. Chelicerae yellow brown; retro- margin with 3 teeth. Legs yellow orange; femur I sometimes black basally; tibiae and tarsi I with fringe of long lateral setae. Abdomen dull yellow brown mesally, darker laterally; venter pale. Terminal apophysis connected to mesal swelling by broad curve; median apophysis with narrow marginal swelling and with distal process weakly hooked (Fig. 14); conductor thin, fluted at tip, lacking knoblike process at tip (Fig. 15). Female. Total length 4.86 ±0.53 mm; carapace 2.40 ±0.17 mm long and 1.89 ±0.15 mm wide (20 specimens). General color and structure as in male, but leg I lacking dark pigment and fringe. Median septum with posterior part approximately rectangular (Fig. 56). Copulatory tubes rather stout, diverging (Fig. 57). Diagnosis. Males of P. mayana most resemble those of P. sagei, P. fastosa, and P. desolatula, but can be distinguished by the narrow marginal swelling and weakly hooked distal process on the median apophysis, and by the broad curve by which the terminal apophysis is connected to the mesal swelling. Females are not anatomically distinguishable from those of sagei, fastosa, and desolatula, but all of those examined were accompanied by males, or originated in localities from which no males of the other species were known. Range. Hidalgo, Mexico to Costa Rica. Natural History. Males and females have been collected in every month except November to January. Females with egg sacs were collected in March and July. Derivation of Specific Name. The name is derived from that of the Maya Indians. 1984] Dondale & Redner — Genus Pardosa 87 Pardosa saxatilis (Hentz) Figures 16, 19-21, 58, 59; Map 3 Lycosa saxatilis Hentz, 1844:392, Figs. 9, 10 (pi. 18). Syntype females from “The mountains of North Alabama”, August, destroyed. Hentz 1875:34, Figs. 9, 10 (pl. 4). Lycosa minima Keyserling, 1877:614, Fig. 3 (pl. 7). Holotype male from Peoria, Illinois, deposited in BM(NH), examined. Banks 1891:193. Pardosa albopatella Emerton, 1885:497, Fig. 2, 2 a, 2b (pl. 49). Four syntype males and 1 juvenile syntype male from New Haven, Connecticut, 20 May 1884 (J. H. Emerton), deposited in MCZ, examined; syntypes from Ipswich and Roxbury, Massachusetts (May and June), not found. Banks 1892:70. Barrows 1918:314. Pardosa annulata Banks, 1892:68, Fig. 41 (pl. 1). Two syntype females from Ithaca, New York, deposited in MCZ, examined. Pardosa minima : Banks 1895:91. Montgomery 1902:571, Figs. 35, 36 (pl. 30); 1904:273. Pardosa saxatilis: Chamberlin 1908:174, Figs. 1, 2 (Pl. 13). Chickering 1933:517. Comstock 1940:661, Fig. 112d. Kaston 1948:335, Fig. 1104 (pl. 56), 1124, 1125 (pl. 58), 1 139 (pl. 59); 1978: 191, Fig. 487 (part). Levi and Field 1954:456. Roewer 1954:194 (part). Bonnet 1958:3420 (part). Wolff 1981:66, Figs. 10, 18. Pardosa platta Chamberlin and Ivie, 1942:31, Fig. 72 (pl. 7). Holotype female from 10 mi. west of Grand Island, Hall Co., Nebraska, 6 June 1933 (W. Ivie), in AMNH, examined. Roewer 1954:194. Vogel 1967: 105. NEW SYNONYM. Male. Total length 4.24 ±0.40 mm; carapace 2.27 ±0.18 mm long and 1.71 ±0.14 mm wide (20 specimens). Carapace (Fig. 20) with dark orange or dark yellow median and submarginal areas, and with paired dark, indistinct, longitudinal bands flanking median area; margins pale or with series of dark spots. Sternum dark orange suffused with black. Chelicerae dark brown, streaked with black; retromargin usually (about 85%) with 2 teeth, more rarely with 3. Legs orange yellow; femur I dark. Abdomen dark orange mottled with brown and black mesally, darker laterally; venter dark orange brown. Terminal apophysis small, blunt; median apophysis long, slender, curved (Fig. 19); conductor sinuous along basal margin, with dark, shiny knob near tip (Fig. 21); patella covered dorsally with reflective white setae, and remaining palpal segments dark (Fig. 16). Female. Total length 4.48 ±0.38 mm; carapace 2.27 ±0.14 mm long and 1.75 ±0.14 mm wide (20 specimens). General color and structure as in male but pale areas on carapace more extensive and more distinct, femur I lacking dark area, rings on distal segments of legs more distinct, and retromargin of chelicera with 2 teeth in 88 Psyche [Vol. 91 approximately 55% of specimens. Median septum extending anteri- ad approximately one-half length of epigynum; hood continuing posteriad at sides where it defines a raised, tapered median area (Fig. 58); copulatory tubes short, slender, with distinct lateral swellings (Fig. 59). Diagnosis. Specimens of P. saxatilis most resemble those of P. atlantica and P. parvula but differ in having the dorsal cover of reflective white setae restricted to the palpal patella. Females are not distinguishable from those of atlantica or parvula except geo- graphically (see map 3). Range. Nebraska and Minnesota to Nova Scotia, south to northern Alabama and North Carolina. No specimens were available from the type locality of P. saxatilis, i.e., northern Alabama, for this study. Thorough collecting there would help to confirm our position regarding the identity of this species. Natural History. Males have been collected in March, May to September, and November, and females May to September. Females with egg sacs were taken in June, July, and September. The usual habitats are grassy fields or meadows but a few specimens also come from marshes, bogs, deciduous woods, or sandy beaches. Wolff (1981) gives life history and related data in Michigan. Pardosa atlantica Emerton Figure 7; Map 3 Pardosa sp. near saxatilis: Banks 1899:189. Pardosa atlantica Emerton, 1913:258, Fig. 7, la (pi. 48). Syntype male and female from Lakehurst, New Jersey, 1 May 1912 (J. H. Emerton), deposited in MCZ, examined; three syntype males from the same locality (same data), deposited in AMNH, examined. Syntype from Fire Island beach, Long Island, New York, not found. Chamberlin and Ivie 1944:147. Pardosa saxatilis: Gertsch 1934:22 (part). Roewer 1954: 194 (part). Bonnet 1958:3420 (part). Pardosa saxatilis var. atlantica: Kaston 1938:184; 1948:335. Howell and Pienkowski 1971:164. Male. Total length 3.53 ±0.27 mm; carapace 1.89 ±0.15 mm long and 1.42 ±0.12 mm wide (20 specimens). Carapace with dark orange or dark yellow median and submarginal areas, and with paired dark, indistinct, longitudinal bands flanking median area; margins 1984] Dondale & Redner — Genus Pardosa 89 pale or with dark spots. Sternum dark orange suffused with black. Chelicerae dark brown, streaked with black; retromargin usually (about 67%) with 3 teeth. Legs orange yellow; femur I dark basally. Abdomen dark orange mottled with brown and black mesally, darker laterally; venter dark orange brown. Terminal apophysis small, blunt; median apophysis long, slender, curved; conductor sinuous along basal margin, with dark, shiny knob near tip; patella, tibia, and small basal area of cymbium covered dorsally with reflective white setae, and remaining palpal segments dark (Fig. 17). Female. Total length 4.03 ±0.50 mm; carapace 1.94 ±0.20 mm long and 1.48 ±0.19 mm wide (20 specimens). General color and structure as in male, but pale areas on carapace more extensive and more distinct, femur I lacking dark area, dark rings on distal segments of legs more distinct, and retromargin of chelicera with 3 teeth in 75% of specimens. Median septum extending anteriad approximately one-half length of epigynum; hood continuing posteriad at sides where it defines a raised, tapered, median area; copulatory tubes short, slender, with distinct lateral swellings. Diagnosis. Males of P. atlantica most resemble those of P. saxatilis and P. parvula but differ in having a dorsal cover of reflective white setae on the patella, tibia, and small basal area of the cymbium of the palpus. Females are not distinguishable from those of saxatilis or parvula except geographically (see map 3). Range. Eastern Oklahoma and eastern Texas to Long Island, New York and Connecticut. The range may extend westward to Kansas, but the westernmost records are based on females, which we are at present unable to distinguish on anatomical characters. Natural History. Males have been collected from February to August, and females from April to August. Females with egg sacs were collected from June to August. The few specimens for which habitat data are available were found in pine or pine-oak forests or at the edge of mixed deciduous forests. One specimen was found in a one-year abandoned field, and Howell and Pienkowski (1971) recorded the species from alfalfa at Blacksburg, Virginia. Emerton’s (1913) type series was collected on “low sandy ground”. Chamberlin and Ivie (1944) found atlantica “in company with "parvula (reported as saxatilis) near Sylvania, Georgia. 90 Psyche [Vol. 91 Pardosa parvula Banks Figure 18; Map 3 Pardosa parvula Banks, 19046:114, Fig. 24 (pi. 6). Holotype male from Altoona, Florida (Dobbin), deposited in MCZ, examined. Banks 1910:59. Pardosa saxatilis: Petrunkevitch 1911:574 (part). Chamberlin and Ivie 1944:149. Muma 1973:180. Male. Total length 3.37 ±0.21 mm; carapace 1.84 ±0.09 mm long and 1.41 ±0.08 mm wide (19 specimens). Carapace with dark orange or dark yellow median and submarginal areas, and with paired dark, longitudinal bands flanking median area; margins pale or dark. Sternum dark orange suffused with black. Chelicerae dark brown, streaked with black; retromargin with 3 teeth. Legs orange yellow; femur I dark basally. Abdomen dark orange mottled with brown and black mesally, darker laterally; venter dark orange brown. Terminal apophysis small, blunt; median apophysis long, slender, curved; conductor sinuous along basal margin, with dark, shiny knob near tip; patella, tibia, and basal half of cymbium covered dorsally with reflective white setae, and remaining palpal segments dark (Fig. 18). Female. Total length 3.84 ±0.43 mm; carapace 1.95 ±0.15 mm long and 1.53 ±0.12 mm wide (20 specimens). General color and structure as in male but pale areas on carapace more extensive and more distinct, femur I lacking dark area, and dark rings on distal segments of legs more distinct. Median septum extending anteriad approximately one-half length of epigynum; hood continuing posteriad at sides where it defines a raised, tapered, median area; copulatory tubes short, slender, with distinct lateral swellings. Diagnosis. Males of P. parvula most resemble those of P. saxatilis and P. atlantica but differ in having a dorsal cover of reflective white setae on the palpal patella, tibia, and basal half of the cymbium. Females are not distinguishable from those of saxatilis and atlantica except geographically (see map 3). Range. Southeastern Louisiana to Florida and southern Georgia. Natural History. Males have been collected from January to April, and in June, July, September, and October, females from January to May, and in August, October, and December. Females with egg sacs were collected in January, March, and December. The 1984] Dondale & Redner — Genus Pardos a 91 recorded habitats are roadside grass and a mowed field, a sugar cane field, and on the ground in goldenrod and pine flat-woods. Chamberlin and Ivie (1944) found parvula (reported as P. saxatilis) “in company with” atlantica near Sylvania, Georgia. Pardosa littoralis Banks Figures 22, 25, 60, 61; Map 1 Pardosa littoralis Banks, 1896:192. Five syntype males, 2 syntype females, and 1 syntype juvenile from Mill Neck, Long Island, New York, June, deposited in MCZ, examined. Emerton 1909:207, Figs. 5, 5a, 5b (pi. 6); 1930:169. Pardosa longispinata Tullgren, 1901:23, Fig. 13 (pi. 1). Holotype female from Lake Leonore, Orange County, Florida (E. Lonnberg), deposited in Zoological Institute, Uppsala University, not examined. Banks 1904a:121; 1910:59. Chamberlin 1908:209. Petrunkevitch 1911:572. Wallace 1950:77, Figs. 3, 4 (pi. 1). Roewer 1954:189. Bonnet 1958:3381. Muma 1973:180; 1975:86. Pardosa floridana Banks, 1904a: 136, Fig. 1 (pi. 7), Fig. 15 (pi. 8). Holotype female from Enterprise, Florida, deposited in MCZ, examined. Banks 1910:59. Gertsch 1934:21. Gertsch and Wallace 1935:5, Figs. 12, 16; 1937:3. Muma 1945:21. Kaston 1948:336, Fig. 1126 (pi. 58), Figs. 1140, 1141 (pi. 59). Bonnet 1958:3369. Pardosa banksi Chamberlin, 1904:175. New name for Pardosa littoralis Banks, mistakenly believed to be preoccupied. Chamberlin 1908:182, Fig. 7 (pi. 13). Banks 1910:58. Petrunkevitch 1911:569. Pardosa ocala Bryant, 1935:81, Fig. 12 (pi. 5). Holotype female from Hale’s Siding, Alachua County, Florida, 14 October 1933 (Wallace), deposited in MCZ, examined. Paratype female from Lake County, 9 October 1933 (H. K. Wallace), not found. Male. Total length 5.07 ±0.42 mm; carapace 2.66 ±0.19 mm long and 2.03 ±0.16 mm wide (20 specimens). Carapace with orange or yellow orange mesal and submarginal areas, and with paired dark, longitudinal bands flanking mesal area; lateral margins usually dark. Sternum orange with marginal black spots and dark mesal band or V-shaped mark, rarely entirely black. Chelicerae orange; retromargin with 3 teeth. Legs orange or yellow orange; femur I sometimes dark basally, usually more so in northern specimens. Abdomen black, sometimes with pale heart mark and series of pale spots at midline, or mottled with yellow brown; heart mark with narrow band of white setae; venter dull yellow, sometimes with small dark spots. Terminal apophysis long, stout, tapered, ex- tending to or beyond tip of embolus; median apophysis broad at base, slender and curved at tip (Fig. 22); conductor with single curve on basal margin, directed retrolaterobasad, with dark, shiny knob near tip (Fig. 25). 92 Psyche [Vol. 91 Female. Total length 5.87 ±0.65 mm; carapace 2.83 ±0.32 mm long and 2.19 ±0.32 mm wide (20 specimens). General color and structure as in male but paler, with the abdominal dorsum having only scattered black spots on a yellowish background. Median septum extending anteriad nearly to level of hood, rather broad anteriorly (Fig. 60); copulatory tubes short, slender, with lateral swellings (Fig. 61). Diagnosis. Specimens of P. littoralis are unique in the milvina group by the greatly elongated terminal apophysis and by the anteriorly broad median septum. Range. Texas to Florida, northward near the coast to the Bay of Fundy, Nova Scotia; Cuba. Natural History. Both sexes of P. littoralis have been collected in every month in the southern part of its range, and egg sacs were observed in April, June to August, and December. In general it is an inhabitant of salt marshes, though Florida records include beaches, lake shores, pine-oak forests, and swamps. One specimen was collected in a soybean field. Muma (1973) trapped many in pine flat- woods, and lesser numbers in a sand-pine dune or in citrus groves. Courtship behavior was described (under the name banksi ) by Kaston (1936). The present authors observed adults running on a salt marsh in Nova Scotia on a warm day at the end of May, 1980. Males and females were seen on or under the mat of dry marsh grasses, particularly at the water’s edge where the mud was wet from the previous tide. If overtaken by a wave they either ran on its surface or climbed emergent plant stems. Comments on Synonymy. This spider has been variously known under five specific names, one of which was proposed on the assumption that the name littoralis was preoccupied in Pardosa. Chamberlin (1904) apparently did not realize that the name Lycosa littoralis Walckenaer, 1805 was a nomen nudum and, therefore, not available. He was apparently also unaware of the older available names Pardosa longispinata Tullgren, 1901 and Pardosa floridana Banks, 1904. Pardosa saltonia sp.n. Figures 23, 26, 62-64; Map 6 Type Material. Holotype male from Fish Springs, west side of Salton Sea, Imperial County, California, 1 16°02'W, 33°25'N, 12 March 1941 (Wilton Ivie), deposited 1984] Dondale & Re drier — Genus Pardos a 93 in AMNH. Two paratype males and 13 paratype females from the type locality (same data as holotype), in AMNH. One paratype female from northeast shore of Salton Sea, 8 October 1963 (Jean and Wilton Ivie), in AMNH. One paratype female from east shore of Salton Sea, 23 September 1957 (Vince Roth), in AMNH. Two paratype males and 1 paratype female from Salton Sea, 26 September 1964 (Vince Roth), in AMNH. One paratype female from Salton, California, in MCZ. Seven paratype males and 5 paratype females from La Choya, Sonora, 12 June 1952 (W. J. Gertsch), in AMNH. Male. Total length 6.39 ±0.35 mm; carapace 3.15 ±0.13 mm long and 2.46 ±0.12 mm wide (12 specimens). Carapace with yellow orange median and submarginal areas, and with pair of dark longitudinal bands flanking median area; margins dark. Sternum orange yellow, suffused with black, or entirely black. Chelicerae orange brown; retromargin with 2 teeth (rarely with small third tooth on one or both chelicerae). Legs dark yellow orange. Abdomen yellow orange mesally, dark laterally; venter pale. Terminal apophysis arched retrolaterally; median apophysis small, occupying only about one-third length of genital bulb (Fig. 23); conductor with single curve on basal margin, with dark, shiny knob near tip (Fig. 26). Female. Total length 7.54 ±1.04 mm; carapace 3.39 ±0.34 mm long and 2.71 ±0.27 mm wide (20 specimens). General color and structure as in male but paler. Epigynum rather short, with ratio of length to greatest width of median septum less than 2:1; septum abruptly narrowed anteriorly from widest point (Fig. 62); copula- tory tubes curved to nearly straight, with swelling on ventral or lateral margin (Figs. 63, 64). Diagnosis. Specimens of P. saltonia differ from those of the other species in the milvina group by the retrolaterally arched terminal apophysis, by the small median apophysis (occupying about one- third the genital bulb length), by the short epigynum and abruptly narrowed median septum (ratio of epigynal length to greatest me- dian septum width less than 2:1), and in part by the presence of 2 teeth on the cheliceral retromargin. Range. Salton Sea area of California; Sonora. Natural History. Males of saltonia have been collected in March, June, and December, and females in March, June, September, October, and December. Egg sacs were found in March and September. 94 Psyche [Vol. 91 Derivation of Specific Name. The name is derived from that of the Salton Sea. Pardosa pauxilla Montgomery Figures 24, 27, 65-67; Map 4 Pardosa pauxilla Montgomery, 1904:268, Figs. 22, 23 (pi. 19). Lectotype male from Austin, Texas, deposited in AMNH and here designated. One paralectotype female from the type locality, in AMNH. One paralectotype male from the type locality, in MCZ. One female, part of the syntype series, deposited in MCZ, represents Pardosa delicatula Gertsch and Wallace. Other syntypes not found. Chamberlin 1908:180, Fig. 9 (pi. 13). Petrunkevitch 1911:574. Gertsch and Wallace 1935:4, Figs. 11, 15. Muma 1945:22; 1973:179; 1975:86. Bonnet 1958:3406. Berry 1970:102. Howell and Pienkowski 1971:164. Pardosa uncatula: Gertsch 1934:20 (part). Roewer 1954:190 (part). Pardosa georgiae Chamberlin and Ivie, 1944:147, Figs. 185, 186 (pi. 13). Holotype female from Savannah Beach, Georgia, 80°51'W, 32°0'N, 4 May 1943 (Wilton Ivie), deposited in AMNH, examined. One paratype male from the type locality, same data, in AMNH, examined. One paratype male and 3 paratype females from the type locality, same data, not found. Wallace 1950:78, Figs. 1, 2 (pi. 1). Roewer 1954:191. Vogel 1967:104. NEW SYNONYM. Male. Total length 4.24 ±0.26 mm; carapace 2.26 ±0.17 mm long and 1.67 ±0.11 mm wide (20 specimens). Carapace usually with yellow orange median and submarginal areas, and with paired dark, longitudinal bands flanking median area; margins usually pale. Sternum dark yellow or yellow brown, sometimes with small black spots along margins. Chelicerae pale yellow brown to black; retromargin with 3 teeth. Legs orange yellow; femur I (and sometimes II) often dark on basal three-fourths. Abdomen dull black, sometimes dull yellow mesally; venter pale yellow, sometimes with faint darker median band. Terminal apophysis with mesal swelling toothlike; median apophysis thick throughout most of its length, with basal process swollen (Fig. 24); conductor with single curve on basal margin, with dark, shiny knob near tip (Fig. 27). Female. Total length 5.05 ±0.55 mm; carapace 2.39 ±0.25 mm long and 1.79 ±0.13 mm wide (20 specimens). General color and structure as in male but paler. Median septum distinctly tapered anteriorly, with expanded posterior part concave at lateral margins (Fig. 65); copulatory tubes rather stout, with swellings on ventral surfaces (Figs. 66, 67). Diagnosis. Specimens of P. pauxilla differ from those of the other species in the milvina group by the toothlike swelling mesal to the 1984] Dondale & Redner — Genus Pardosa 95 terminal apophysis, by the large, swollen basal process on the median apophysis, and by the long, distinctly tapered median septum of which the posterior expanded part has concave lateral margins. Range. Kansas to New Jersey, south to New Mexico, southern Texas, and Florida. Natural History. Males and females have been collected in every month except December. Egg sacs were collected from March to May and from July to November. The commonest habitat is grassy pastures, but many specimens have been taken at stream or pond margins or in crops such as alfalfa, soybeans, sweet potatoes, and peanuts. Muma (1973) trapped specimens in citrus groves and pine flat-woods in Florida. Pardosa portorieensis Banks Figures 28, 29, 68, 69 Pardosa porto-ricensis Banks, 1902:224, Figs. 2, 3 (pi. 15). Holotype female from San Juan, Puerto Rico, 1-10 January 1899 (August Busck), deposited in USNM, examined. Pardosa portorieensis: Petrunkevitch 1911:574; 1929:87, Figs. 74, 75. Roewer 1954:188. Bonnet 1958:3407. Male. Total length 3.56 mm; carapace 2.00 mm long and 1.56 mm wide (1 specimen). Carapace with orange yellow median and submarginal areas, and with pair of distinct dark, longitudinal bands flanking median area; margins somewhat dark. Sternum pale yellow. Chelicerae pale orange, lightly marked with gray; retro- margin with 3 teeth. Legs pale yellow. Abdomen pale mesally, with pair of dark longitudinal bands laterally. Terminal apophysis small, pointed; median apophysis with distal process expanded (ventral view, Fig. 28); conductor curved on basal margin, with dark shiny knob near tip (Fig. 29). Female. Total length 4.51 ±0.70 mm; carapace 2.24 ±0.26 mm long and 1.79 ±0.23 mm wide (16 specimens). General color and structure as in male. Epigynum rather long (ratio of epigynum length to greatest septal width more than 2:1) (Fig. 68); median septum tapered and indistinct anteriorly; copulatory tubes slender, and spermathecae narrowly separated (Fig. 69). 96 Psyche [Vol. 91 Map 4. Collection localities for Pardosa pauxilla. Map 5. Collection localities for Pardosa milvina. 1984] Dondale & Redner — Genus Pardosa 97 Diagnosis. Specimens of P. portoricensis differ from those of the other West Indian species in the milvina group by the expanded distal process of the median apophysis, by the longitudinally banded abdominal dorsum, by the long epigynum (ratio of epigynal length to greatest septal width more than 2:1), by the anteriorly tapered median septum, and by the slender copulatory tubes and narrowly separated spermathecae. Localities. Puerto rico: San Juan; Mayaguez; Martin Pena; Puerto Nuevo Point, near Vega Baja; Embarcadero Point, near Luquillo. virgin islands: St. Croix: Chistiansted. antigua: Lignum Vitae Bay: Reed Point, Jolly Beach. Range. Puerto Rico, Virgin Islands, and western Antigua. Natural History. The only available male was collected with females in October. Other females were collected in January to March and in June. Egg sacs were found in June, and one of the October females carried young. Petrunkevitch (1929) collected specimens on the “sandy plains” of the northern coast of Puerto Rico. Pardosa milvina (Hentz) Figures 30-32, 70, 71; Map 5 Lycosa milvina Hentz, 1844:392, Fig. 8 (pi. 18). Holotype female from Alabama, September, destroyed. Hentz 1875:33, Fig. 8 (pi. 4). Lycosa flavipes Keyserling, 1877:616, Fig. 4 (pi. 7). Syntype female from Peoria, Illinois, deposited in BM(NH), examined. Syntype female from Baltimore, Maryland, not found. Pardosa nigropalpis Emerton, 1885:497, Figs. 1, 1 a-ld (pi. 49). Five syntype males and 1 syntype female from New Haven, Connecticut, 25 May 1883 (J. H. Emerton), deposited in MCZ, examined. Syntypes from Massachusetts, not found. Emerton 1902:83, Figs. 202-204. Stone 1890:430. Banks 1892:70; 1893:125. Montgomery 1902:569, Figs. 32-34 (pi. 30); 1904:275. Scheffer 1905a: 119; 19056:191. Bryant 1908:89. Barrows 1918:315. Roewer 1954:193. Pardosa milvina: Banks 1899:189; 1904a: 135; 19046:115, Fig. 6 (pi. 5). Scheffer 1906:126. Chamberlin 1908:177, Figs. 3, 4 (pi. 13). Petrunkevitch 1911:572. Gertsch and Wallace 1935:5, Figs. 19, 20. Comstock 1940:662, Figs. 731/', 732e. Muma 1945:22. Kaston 1948:334, Figs. 1100-1103 (pi. 56), 1138 (pi. 59); 1978:190, Fig. 486. Levi and Field 1954:456. Roewer 1954:192. Bonnet 1958:3388. Fitch 1963:113. Berry 1970:102. Howell and Pienkowski 1971:164. Wolff 1981:64. 98 Psyche [Vol. 91 Pardosa scita Montgomery, 1902:573, Figs. 37, 38 (pi. 30). One syntype male and 1 syntype female from Philadelphia, Pennsylvania, deposited in AMNH, exam- ined. Montgomery 1904:272. Pardosa canadensis: Banks 1910:58 (part); 1916:81 (part). Bonnet 1958:3362 (part). Male. Total length 4.64 ±0.36 mm; carapace 2.40 ±0.20 mm long and 1.91 ±0.15 mm wide (20 specimens). Carapace with yellow orange median and submarginal areas, and with paired black longitudinal bands flanking median area; margins often pale. Sternum yellow to nearly black. Chelicerae pale to dark yellow brown; retromargin with 3 teeth. Legs orange yellow; femur I sometimes black on basal half. Abdomen yellow brown, much mottled with black; venter pale yellow, often lightly spotted or lined with black. Terminal apophysis broad, blunt; median apophysis slender at middle, with distal process slender (Figs. 30, 31); conductor curved on basal margin, with dark, shiny knob near tip (Fig. 32). Female. Total length 5.77 ±0.65 mm; carapace 2.73 ±0.25 mm long and 2.16 ±0.19 mm wide (20 specimens). General color and structure as in male but legs more distinctly ringed and abdominal dorsum sometimes with paried, indistinct longitudinal bands; femur I lacking black area in basal half. Median septum extending anteriad less than one-half length of epigynum; hood continuing posteriad at sides where it defines a depressed, non-tapered area (Fig. 70); copulatory tubes rather slender, with lateral swellings (Fig. 71). Diagnosis. Specimens of P. milvina are distinguished from those of the other species in the milvina group by the combination of terminal apophysis small, broad, blunt, arched mesally, and lacking a toothlike mesal swelling, median apophysis slender at middle and with a slender distal process, short median septum, hood extending posteriad at sides where it defines a depressed, non-tapered area, and occurrence only north of the Tropic of Cancer (approximately 23°30'N). Range. Northern Peninsula of Michigan and southern Ontario to southern Quebec and Maine, south to Texas and Florida. Natural History. Males of P. milvina have been collected from February to August, and females from February to November. Egg sacs were recorded from April to September. The species appears to 1984] Dondale & Redner — Genus Pardosa 99 reach high densities in moist habitats such as swamps, meadows, mud flats, and edges of creeks and ponds, but is also found in deciduous and cedar woods, lawns, gardens, pastures, and various crops (rice, corn, cotton, sweet potato, soybean, alfalfa). Mont- gomery (1903) and Kaston (1936) described courtship and copula- tory behavior. Pardosa guadalajarana sp.n. Figures 33-35, 72, 73; Map 6 Type Material. Holotype male from 4 miles southwest of Guada- lajara, Jalisco, Mexico, 20 June 1941 (L. I. Davis), deposited in AMNH. Three paratype males and three paratype females from the type locality (same data as holotype), deposited in AMNH. One paratype male and fourteen paratype females from localities other than the type locality, in Mexico and Honduras, deposited in AMNH. Male. Total length 3.03-3.18 mm; carapace 1.62-1.70 mm long and 1.21-1.30 mm wide (5 specimens). Carapace black, with dark orange brown median area. Sternum dark orange brown suffused with black. Chelicerae black, with dull yellow area anteromesally; retromargin with 2 teeth. Legs orange brown, with femora often darkened or with 2 or 3 dark, indistinct rings. Abdomen dark red brown, spotted with black, and with median band of reflective white setae; venter dull yellow, sometimes with small black spots. Terminal apophysis pointed; median apophysis abruptly angled on mesal margin (Figs. 33, 35); conductor with single curve along basal margin, with dark knob near tip (Fig. 34). Female. Total length 3.96 ±0.36 mm; carapace 1.86 ±0.08 mm long and 1.42 ±0.11 mm wide (16 specimens). General color and structure as in male but carapace usually much paler, the carapace with pale median band distinct and margins pale, and abdomen lacking band of reflective white setae. Median septum extending approximately three-fourths length of epigynum; posterior part more or less triangular, concave at lateral margins, and rather narrow (ratio of epigynal length to greatest septum width more than 2:1) (Fig. 72); copulatory tubes slender, with lateral swellings (Fig. 73). 100 Psyche [Vol. 91 Diagnosis. Specimens of P. guadalajarana most resemble those of P. longivulva and P. marialuisae but differ in the abruptly angled median apophysis, in the possession of two retromarginal teeth on the chelicera, and in the small size. Range. Jalisco to the Honduras/ El Salvador border. Natural History. Males have been collected in June and July, and females June to August. No habitat data are recorded. Derivation of Specific Name. The name is derived from that of the city of Guadalajara. Pardosa longivulva F. Pickard-Cambridge Figures 36-38, 74, 75; Map 6 Pardosa longivulva F. Pickard-Cambridge, 1902:318, Figs. 21, 21a, 22 (pi. 30). Holotype female from Teapa, Tabasco, Mexico (H. H. Smith), deposited in BM(NH), examined. Two paratype males from the type locality (same vial as holotype), in the same institution, examined. One female from the type locality (same vial as holotype and paratypes) is Pardosa hamifera F. Pickard- Cambridge. Paratype from Amula, Guerrero, Mexico, not found. Petrunkevitch 1911:572. Pardosa longivulvula Roewer, 1951:440. New name for Pardosa longivulva, proposed on the mistaken assumption that the latter was preoccupied. Male. Total length 4.31-4.50 mm; carapace 2.08-2.49 mm long and 1.72-2.03 mm wide (6 specimens). Carapace with orange median and submarginal areas, and with paired dark, longitudinal bands flanking median area; margins black. Sternum dark orange, often with scattered black spots or with median black band. Chelicerae dark orange brown to black; retromargin with 3 teeth. Legs yellow orange; femur I sometimes with black area at base; tibia and basitarsus I with fringe of long, erect, dark setae on prolateral and retrolateral surfaces. Abdomen black to yellow brown, darker laterally; venter off-white to yellow brown, sometimes with many dark dots or with one to three dark longitudinal bands. Terminal apophysis long, rather slender; median apophysis slender at middle, with slender distal process (Figs. 36, 37); conductor with single curve along basal margin, with dark knob near tip (Fig. 38). Female. Total length 4.98-5.47 mm; carapace 2.58-2.80 mm long and 1.96-2.42 mm wide (4 specimens). General color and structure 1984] Dondale & Redner — Genus Pardosa 101 as in male but carapace with pale areas more extensive and leg I lacking dark area at base and lacking setal fringe. Epigynum rather long (ratio of epigynal length to greatest septal width more than 2:1); median septum with expanded posterior part approximately triangular in outline and concave at lateral margins (Fig. 74); copulatory tubes with small swellings at base (Fig. 75). Diagnosis. Specimens of P. longivulva can be distinguished from those of the other species in the milvina group by the combination of black carapace margins, presence of a setal fringe on tibia and basitarsus I of males, rather long and slender terminal apophysis, long epigynum (ratio of epigynal length to greatest septal width more than 2:1), small swellings at base of copulatory tubes, and occurrence restricted south of the Tropic of Cancer. Range. Tabasco, Mexico to Guatemala. Natural History. Adults have been collected in February, March, and June. One of the females collected in February had an egg sac. Habitat is not recorded. Pardosa marialuisae sp.n. Figures 39, 40, 42, 76, 77; Map 6 Type Material. Holotype male from Pico de Oro, 93°30'W, 17°58'N, Tabasco, Mexico, 12 August 1966 (Jean and Wilton Ivie), deposited in AMNH. Paratype female from the type locality (same data as holotype), in AMNH. Twenty-four paratype males and 33 paratype females from Mexico and Honduras, deposited in AMNH. Five paratype males and 17 paratype females from Guatemala, deposited in MCZ. Male. Total length 3.85 ±0.20 mm; carapace 2.04 ±0.10 mm long and 1.57 ±0.12 mm wide (20 specimens). Carapace usually black, with margins alone pale, but rarely with pale submarginal areas and dark margins. Sternum orange yellow, sometimes with small black spots at posterior and lateral margins. Chelicerae usually black, more rarely yellow orange; retromargin with 3 teeth. Legs yellow orange; femur I sometimes with black area at base. Abdomen yellow orange to black; if pale, then mottled or spotted, and darkest along lateral margins; venter pale yellow, sometimes spotted with black. 102 Psyche [Vol. 91 Map 6. Collection localities for Pardosa saltonia (triangles), P. quadalajarana (half-circles), P. longivulva (stars), marialuisae (circles). 1984] Dondale & Redner — Genus Pardosa 103 Terminal apophysis slender, pointed, rather short; median ap- ophysis slender at middle, with slender distal process (Figs. 39, 42); conductor curved on basal margin, with dark, shiny knob near tip (Fig. 40). Female. Total length 4.80 ±0.64 mm; carapace 2.30 ±0.18 mm long and 1.88 ±0.19 mm wide (20 specimens). General color and structure as in male but with pale mesal area on carapace, and with front and chelicerae pale. Epigynum long (ratio of total length to greatest median septum width more than 2:1); median septum with expanded posterior part approximately triangular, concave at sides (Fig. 76); copulatory tubes with large swellings at lateral margins (Fig. 77). Diagnosis. Specimens of P. marialuisae can be distinguished from the other members of the milvina group by the combination of usually pale carapace margins, lack of a setal fringe on leg I, the possession of a rather short, pointed terminal apophysis, long epigynum (ratio of epigynal length to greatest septal width more than 2:1), large swellings at the sides of the copulatory tubes, and occurrence restricted south of the Tropic of Cancer. Range. Hidalgo, Mexico to Honduras. Natural History. Males have been collected from February to August and in October, females from January to August and in October. Egg sacs were collected in May and August. The only habitat recorded on the labels is “along river” at 30 miles southeast of Palenque Aqua Azul, Chiapas, Mexico. Derivation of Specific Name. The name is derived from that of our friend and colleague Maria-Luisa Jimenez of Universidad Nacional Autonoma de Mexico. Acknowledgments The authors are deeply indebted to the following who lent specimens for this study: Mr. J. C. Cokendolpher, Texas Tech University; Dr. G. B. Edwards, Florida State Collection of Arthropods; Dr. J. S. Heiss, University of Arkansas; Mr. G. F. Hevel, United States National Museum (USNM); Dr. N. Horner, Midwestern State University; M. en C. Maria-Luisa Jimenez, Universidad Nacional Autonoma de Mexico; Drs. J. Kekenbosch 104 Psyche [Vol. 91 and L. Baert, Institut Royal des Sciences Naturelles de Belgique; Dr. H. W. Levi, Museum of Comparative Zoology, Harvard University (MCZ); Dr. Patricia R. Miller, Mississippi Entomo- logical Museum; Mr. A. J. Penniman, Ohio State University; Dr. N. I. Platnick, American Museum of Natural History (AMNH); Dr. C. L. Remington, Peabody Museum of Natural History; Mr. F. R. Wanless and Mr. P. D. Hillyard, British Museum (Natural History) (BMNH). Literature Cited Banks, N. 1891 Synonymical notes in spiders. Ent. News 2:193. Banks, N. 1892 The spider fauna of the upper Cayuga Lake basin. Proc. Acad. Nat. Sci. Philad. 1892:11-81. Banks, N. 1893 Notes on spiders. J. N.Y. Ent. Soc. 1:123-134. Banks, N. 1895 A list of spiders of Long Island, with description of new species. J. N.Y. Ent. Soc. 3:76-93. Banks, N. 1896 Additions to the list of Long Island spiders. J. N.Y. Ent. Soc. 4:190-192. Banks, N. 1898 Arachnida from Baja California, and other parts of Mexico. Proc. Cal. Acad. Sci. (ser. 3) 1:205-308. Banks, N. 1899 Some spiders from northern Louisiana. Proc. Ent. Soc. Wash. 4: 188-195. Banks, N. 1902 Some spiders and other Arachnida from Porto Rico. Proc. U.S. Nat. Mus. 24:217-227. Banks, N. 1904a The Arachnida of Florida. Proc. Acad. Nat. Sci. Philad. 56:120-147. Banks, N. 1904b New genera and species of Nearctic spiders. J. N.Y. Ent. Soc. 12:109-119. Banks, N. 1909 Arachnida from Costa Rica. Proc. Acad. Nat. Sci. Philad. 61:194-234. Banks, N. 1910 Catalogue of Nearctic spiders. Bull. U.S. Nat. Mus. 72:1-80. Banks, N. 1916 Revision of Cayuga Lake spiders. Proc. Acad. Nat. Sci. Philad. 68:68-84. Barrows, W. M. 1918 A list of Ohio spiders. Ohio J. Sci. 18:297-318. 1984] Dondale & Redner — Genus Pardosa 105 Berry, J. W. 1970 Spiders of the North Carolina Piedmont old-field communities. J. Elisha Mitchell Sci. Soc. 86:97-105. Bonnet, P. 1958 Bibliographia Araneorum. Tome 2, pt. 4. Imprimerie Douladoure, Toulouse. Bryant, E. B. 1908 List of the Araneina. In Fauna of New England, 9. Occ. Pap. Boston Soc. Nat. Hist. 7:1-105. Bryant, E. B. 1935 A few southern spiders. Psyche 42:73-83. Chamberlin, R. V. 1904 Notes on generic characters in the Lycosidae. Can. Ent. 36:145-148, 173-178. Chamberlin, R. V. 1908 Revision of North American spiders of the family Lycosidae. Proc. Acad. Nat. Sci. Philad. 60:158-318. Chamberlin, R. V. and W. Ivie 1942 A hundred new species of American spiders. Bull. Univ. Utah. (Biol. Ser.) 32(13): 1-1 17. Chamberlin, R. V. and W. Ivie 1944 Spiders of the Georgia region of North America. Bull. Univ. Utah (Biol. Ser.) 35(9): 1-267. Chickering, A. M. 1933 Notes and studies on Arachnida. IV. Araneae from the Douglas Lake region, Michigan II. Pap. Mich. Acad. Sci. 17:515-520. Comstock, J. H. 1940 The spider book (rev. and ed. by W. J. Gertsch). Comstock Publishing Company, Inc., Ithaca. Dondale, C. D. and J. H. Redner 1978 Revision of the Nearctic wolf spider genus Schizocosa (Araneida: Lycosidae). Can. Ent. 110:143-181. Emerton, J. H. 1885 New England Lycosidae. Trans. Conn. Acad. Arts Sci. 6:481-505. Emerton, J. H. 1909 Supplement to the New England spiders. Trans. Conn. Acad. Arts Sci. 14:171-236. Emerton, J. H. 1913 New and rare spiders from within fifty miles of New York City. Bull. Amer. Mus. Nat. Hist. 32:255-260. Emerton, J. H. 1930 Spiders of Nantucket. Publ. Nantucket Maria Mitchell Assoc. 3: 161-174. Fitch, H. S. 1963 Spiders of the University of Kansas Natural History Reservation and Rockefeller Experimental Tract. Misc. Publ. Univ. Kansas, No. 33, pp. 1-202. Gertsch, W. J. 1934 Notes on American Lycosidae. Amer. Mus. Novit., No. 693, pp. 1-25. 106 Psyche [Vol. 91 Gertsch, W. J. and L. I. Davis 1940 Report on a collection of spiders from Mexico II. Amer. Mus. Novit., No. 1059, pp. 1-18. Gertsch, W. J. and H. K. Wallace 1935 Further notes on American Lycosidae. Amer. Mus. Novit., No. 794, pp. 1-22. Gertsch, W. J. and H. K. Wallace 1937 New American Lycosidae with notes on other species. Amer. Mus. Novit., No. 919, pp. 1-22. Hentz, N. M. 1844 Descriptions and figures of the Araneides of the United States. Boston J. Nat. Hist. 4:386-396. Hentz, N. M. 1875 The spiders of the United States, a collection of the arachnological writings of Nicholas Marcellus Hentz, M. D., edited by Edward Burgess with notes and descriptions by James H. Emerton. Occ. Pap. Boston Soc. Nat. Hist. 2:1-171. Howell, J. O. and R. L. Pienkowski 1971 Spider populations in alfalfa, with notes on spider prey and effect of harvest. J. Econ. Ent. 64:163-168. Kaston, B. J. 1936 The senses involved in the courtship of some vagabond spiders. Ent. Amer. 16:97-167. Kaston, B. J. 1938 Checklist of the spiders of Connecticut. Bull Conn. Geol. Nat. Hist. Surv. 60:175-201. Kaston, B. J. 1948 Spiders of Connecticut. Bull. Conn. Geol. Nat. Hist. Surv. 70:1-874. Kaston, B. J. 1978 How to know the spiders. 3rd Ed. Wm.C. Brown Company Publishers, Dubuque. Keyserling, E. 1877 Ueber amerikanische Spinnenarten der Unterordnung Citigradae. Verh. Zool. Bot. Ges. Wien 26:609-708. Levi, H. W. and H. M. Field 1954 The spiders of Wisconsin. Amer. Midi. Nat. 51:440-467. Lowrie, D. C. and C. D. Dondale 1981 A revision of the nigra group of the genus Pardosa in North America (Araneae, Lycosidae). Bull. Amer. Mus. Nat. Hist. 170:125-139. Montgomery, T. H., Jr. 1902 Descriptions of Lycosidae and Oxyopidae of Philadelphia and its vicinity. Proc. Acad. Nat. Sci. Philad. 54:534-592. Montgomery, T. H., Jr. 1903 Studies on the habits of spiders, particularly those of the mating period. Proc. Acad. Nat. Sci. Philad. 55:59-149. Montgomery, T. H., Jr. 1904 Descriptions of North American Araneae of the families Lycosidae and Pisauridae. Proc. Acad. Nat. Sci. Philad. 56:261-323. 1984] Dondale & Redner — Genus Pardosa 107 Muma, M. H. 1945 An annotated list of the spiders of Maryland. Bull. Univ. Maryland Agr. Expt. Sta., A38, pp. 1-65. Muma, M. H. 1973 Comparison of ground surface spiders in four central Florida eco- systems. Fla. Ent. 56:173-196. Muma, M. H. 1975 Spiders in Florida citrus groves. Fla. Ent. 58:83-90. Petrunkevitch, A. 1911 A synonymic index-catalogue of spiders of North, Central and South America. ..Bull. Amer. Mus. Nat. Hist. 29:1-791. Petrunkevitch, A. 1925 Arachnida from Panama. Trans. Conn. Acad. Arts Sci. 27:51-248. Petrunkevitch, A. 1929 The spiders of Porto Rico. Part one. Trans. Conn. Acad. Arts Sci. 30:1-158. Pickard-Cambridge, F. 1902 Arachnida. Araneida 2. Biol. Cent. -Amer., Zool., pp. 313-424. Roewer, C. F. 1951 Neue Namen einiger Araneen-Arten. Abh. Naturw. Ver. Bremen 32:437-456. Roewer, C. F. 1954 Katalog der Araneae. Bd. 2. Institut Royal des Sciences Naturelles de Belgique, Bruxelles. Scheffer, T. H. 1905a List of spiders in the entomological collection of the Kansas State University. Bull. Kans. St. Univ. 3:115-120. Scheffer, T. H. 1905b A preliminary list of Kansas spiders. Trans. Kans. Acad. Sci. 19: 182-193. Scheffer, T. H. 1906 Additions to the list of Kansas Arachnida. Trans. Kans. Acad. Sci. 20:121-130. Stone, W. 1890 Pennsylvania and New Jersey spiders of the family Lycosidae. Proc. Acad. Nat. Sci. Philad. 1890:420-434. Tullgren, A. 1901 On the spiders collected in Florida by Dr. Einar Lonnberg 1892-93. Bih. Svenska Vet.-Akad. Handl. 27:1-29. Vogel, B. R. 1967 A list of new North American spiders 1940-1966. Mem. Amer. Ent. Soc., No. 23, pp. 1-186. Wallace, H. K. 1950 On Tullgren’s Florida spiders. Fla. Ent. 33:71-83. Wolff, R. J. 1981 Wolf spiders of the genus Pardosa (Araneae: Lycosidae) in Michigan. Gt. Lakes Ent. 14:63-68. 108 Psyche [Vol. 91 Figs. 2-7. External male genitalia of Pardosa spp. 2-4, palpus, ventral view. 5-7, apical division, ventral view. 2,5, P. bellona Banks. 3,6, P. delicatula Gertsch and Wallace. 4,7, P. hamifera F. Pickard-Cambridge. bp, basal process of median apophysis, con, conductor, cym, cymbium. dp, distal process of median apophysis, e, embolus, ma, median apophysis./?, palea. teg, tegulum. term, terminal apophysis. 1984] Dondale & Redner — Genus Pardosa 109 Figs. 8-15. External male genitalia of Pardosa spp. 8-10,14, palpus, ventral view. 1 1-13,15, apical division, ventral view. 8,1 1, P. sagei Gertsch and Wallace. 9,12, P. fastosa (Keyserling). 10,13, P. desolatula Gertsch and Davis. 14,15, P. may ana sp.n. [Vol. 91 Figs. 16-21. Male structures of Pardosa spp. 16-18, palpus, dorsal view. 19, palpus, ventral view. 20, body, dorsal view. 21, apical division, ventral view. 16,19-21, P. saxatilis (Hentz). 17, P. atlantica Emerton. 18, P. parvula Banks. 1984] Dondale & Re drier — Genus Pardosa 111 Figs. 22-29. External male genitalia of Pardosa spp. 22-24, 28, palpus, ventral view. 25-27,29, apical division, ventral view. 22,25, P. littoralis Banks. 23,26, P. saltonia sp. n. 24,27, P. pauxilla Montgomery. 28,29, P. portoricensis Banks. 112 Psyche [Vol. 91 Figs. 30-38. External male genitalia of Pardosa spp. 30,33,36, apical division, retrolateral view. 31,35,37, palpus, ventral view. 32,34,38, apical division, ventral view. 30-32, P. milvina (Hentz). 33-35, P. guadalajarana sp. n. 36-38, P. longivulva F. Pickard-Cambridge. 1984] Dondale & Redner — Genus Pardosa 113 Figs. 39-45. External male genitalia of Pardosa spp. 39, apical division, retrolateral view. 40, apical division, ventral view. 42, male palpus, ventral view. 41,43-45, epigynums. 41, ventral view. 43,44, dorsal view. 45, lateral view. 39,40,42, P. marialuisae sp. n. 41, 43, 44, P. bellona Banks. 45, P. delicatula Gertsch and Wallace, co, copulatory opening, ct, copulatory tube, h, hood. ms. median septum, spt, spermathecae. Figs. 46-53. Epigynums of Pardosa spp. 46,48,50,52, ventral view. 47,49,51,53, dorsal view. 46,47, P. delicatula Gertsch and Wallace. 48,49, P. hamifera F. Pickard-Cambridge. 50, 51, P. sagei Gertsch and Wallace. 52,53, P. fastosa (Keyserling). 1984] Dondale & Redner — Genus Pardosa 115 Figs. 54-61. Epigynums of Pardosa spp. 54,56,58,60, ventral view. 55,57,59,61, dorsal view. 54,55, P. desolatula Gertsch and Davis. 56,57, P. mayana sp. n. 58, 59, P. saxatilis (Hentz). 60,61, P. littoralis Banks. 116 Psyche [Vol. 91 Figs. 62-69. Epigynums of Pardosa spp. 62,65,68, ventral view. 63,64,66,67,69, dorsal view. 62-64, P. saltoma sp 65-67, P. pauxilla Montgomery. 68, 69, P. portoricensis Banks. Figs. 70-77. Epigynums of Pardosa spp. 70,72,74,76, ventral view. 71,73,75,77, dorsal view. 70,71, P. milvina (Hentz). 72,73, P . guadalajarana sp. n. 74,75, P. longivulva F. Pickard-Cambridge. 76,77, P. marialuisae sp. n. THE IDENTITY OF CHAETOCLUSIA AFFINIS JOHNSON AND ITS PLACEMENT IN SOBAROCEPHALA CZERNY (DIPTERA: CLUSIIDAE)* By Norman E. Woodley Systematic Entomology Laboratory, IIBIII, ARS, USDA c/o U. S. National Museum of Natural History Washington, DC 20560 Johnson (1913) described Chaetoclusia affinis on the basis of one male and one female from New Jersey. His brief and inadequate description has not allowed subsequent authors to recognize the species and omits discussion of characters later used to define clusiid genera. Melander and Argo (1924) simply repeated Johnson’s description and mentioned that they had not seen his material. I have recently examined the holotype male and allotype female of C. affinis, which are housed in the Museum of Comparative Zoology at Harvard University (MCZ #7873). It was readily apparent that Johnson had placed the species in the wrong genus, and that C. affinis actually belongs in the genus Sobarocephala Czerny, not Chaetoclusia Coquillett. Both of Johnson’s specimens lack the longer, bristle-like hairs on wing vein R,, which are characteristic of Chaetoclusia. Johnson probably placed the species in Chaetoclusia because of its densely pubescent antennal arista, a character state found in Chaetoclusia bakeri Coquillett, the type species of the genus. This feature, however, is not unique to Chaetoclusia, and in fact partially defines the “plumata” species group of Sobarocephala, as recognized by So6s (1963). When S. affinis is run through the most recent key to Nearctic Sobarocephala (Sabrosky and Steyskal, 1974), it runs without difficulty to S. testacea So6s. The holotype male of S. testacea, housed in the U. S. National Museum of Natural History, was recently examined and found to be conspecific with S. affinis. Johnson’s allotype is probably also conspecific, but there is difficulty in separating females of this species from those of S. muesebecki Sabrosky and Steyskal. The latter species is known with * Manuscript received by the editor February 15, 1984. 119 120 Psyche [Vol. 91 certainty from Florida, and from a questionable specimen from Maryland (see Sabrosky and Steyskal, 1974: 378). As a result of this study, the following nomenclatural changes result: Sobarocephala affinis (Johnson), 1913: 101. new combination. Sobarocephala testacea So6s, 1964: 449. new synonymy. The holotype of Chaetoclusia affinis Johnson is in poor condition. It is missing the head and the legs beyond the trochanters, except that the right front femur and tibia are present. I have removed and cleared most of the abdomen, including the male terminalia, and placed them in glycerin in a microvial on the specimen pin, so that they will be better preserved and more useful for future workers. Due to the absence of the head, the male genitalia are the only features of the type that will allow conclusive identification of the specimen. Because C. affinis Johnson really belongs in Sobarocephala, this deletes the only species ascribed to Chaetoclusia from the Nearctic fauna. The genus is now known only from the Neotropical Region extending from Nicaragua south to Peru, and also Haiti and St. Vincent (So6s, 1968). It seems unlikely that the genus will be found in the Nearctic Region, although its Caribbean distribution makes it just possible that it might someday be found in southern Florida. I wish to thank Wayne Mathis, Department of Entomology, Smithsonian Institution, and D. R. Miller, Systematic Entomology Lab, USDA, for reviewing the manuscript. Literature Cited Johnson, C. W. 1913. A study of the Clusiodidae, (Heteroneuridae) of the eastern United States. Psyche 20:97-101. Melander, A. L. and N. G. Argo. 1924. A revision of the two-winged flies of the family Clusiidae. Proc. U. S. Nat. Mus. 64:1-54. Sabrosky, C. W. and G. C. Steyskal. 1974. The genus Sobarocephala (Diptera: Clusiidae) in America north of Mexico. Ann. Ent. Soc. Amer. 67:371-385. So6s, A. 1963. Identification key to the species of the "‘plumata- group” of the genus Sobarocephala Czerny (Diptera: Clusiidae). Acta Zool. Acad. Sci. Hung. 9:391-396. 1984] Woodley — Chaetoclusia 121 1964. New Sobarocephala-species from the "plumata- group” (Diptera, Clu- siidae). Ann. Hist.-nat. Mus. Nat. Hung. 56:449-455. 1968. Family Clusiidae, No. 84. In A catalogue of the Diptera of the Americas south of the United States. SSo Paulo, Museu de Zoologia, Universidade de Sao Paulo, 13 pp. THE INFLUENCE OF MICROHABITAT AND PREY AVAILABILITY ON BURROW ESTABLISHMENT OF YOUNG GEOLYCOSA TURRICOLA (TREAT) AND G. MICANOPY WALLACE (ARANEAE: LYCOSIDAE): A LABORATORY STUDY* By G. L. Miller Department of Entomology, Mississippi Entomological Museum, Drawer EM, Mississippi State University Mississippi State, MS 39762 Introduction The survival of an animal largely depends on its ability to locate and use a suitable habitat. The suitability of a habitat will depend on such things as prey availability, microhabitat characteristics and the interaction of these, but our understanding of the interaction of these factors and how they effect the animal’s choice of habitat is poor (Krebs, 1978). For spiders, there have been many studies which show the importance of habitat characteristics and prey abundance in the selection of foraging and web sites (e.g., Savory, 1930; Enders, 1977; Riechert, 1976; Riechert and Tracy, 1975). However, most studies deal with adult spiders (some exceptions being the work of Waldorf, 1976; Enders, 1977 and Hallander, 1970) and our understanding of which factors may influence habitat selection in newly dispersing young spiders is limited. These factors are particularly important for burrowing wolf spiders ( Geolycosa ) since the selected burrow site is generally used throughout the life of the spider (Wallace, 1942). In this paper I test selected hypotheses about the interactions among burrow establishment, prey availability, and several microhabitat characteristics in two species of burrowing wolf spiders. This paper is not concerned with the relationship between the burrow site characteristics and survival of the spiderling, which is best studied in the field (see Reichert, 1976 and Reichert and Tracy, 1975 for * Manuscript received by the editor January 18, 1984. 123 124 Psyche [Vol. 91 examples of elegant field studies concerning habitat selection and survival). Geolycosa spp. are obligate burrowers which establish a burrow shortly after leaving the mother (Wallace, 1942; McCrone, 1964; pers. observ.) and, with the exception of short foraging sorties and the reproductive wanderings of mature males, live their entire lives within a burrow (Wallace, 1942; McCrone, 1964; McQueen, 1978; Humphreys, 1975; pers. observ.). Generally, newly-dispersing spi- derlings construct burrows in the vicinity of the maternal burrow (McQueen, 1978). Methods and Materials Specimens — I collected Geolycosa micanopy from Alachua, Levy, Marion and Putnam counties Florida during December 1982 and March 1983 and G. turricola from Oktibbeha County, Mississippi in March 1983. The spiders were housed at room temperature (21° C) and lighting (ca. 10:14 light:dark) in individual translucent plastic cups containing sand. Most of the spiders constructed burrows in the cups. The spider’s diet consisted of crickets, wingless flies and tobacco budworm larvae (Heliothis sp.); and free water was provided. Several spiders had mated before collection and constructed egg cases in the lab. Most females held the egg case until the young emerged and then tolerated them on her back until they dispersed. However, two G. micanopy and one G. turricola cast the cases from the burrows. The young from these cases were used in the experiments because their feeding experience could be closely controlled. Abandoned cases were kept in individual petri dishes. After about two weeks, I made a small opening in the egg sac and the young emerged. The number of young used for the study was 121 and 98 for the two G. micanopy egg cases and 119 for the G. turricola egg case. Spiderlings congregated on a cotton ball placed near the egg case. Each individual was placed into a glass vial within one week of emergence and held without food (water provided via cotton swab) for about two weeks, at which time the experiments began. Experimental design and hypotheses — Three experiments were designed to test hypotheses concerning independence among dichotomous grouping variables arranged in three three-way 1984] Miller — Geolycosa 125 Table 1. Observed cell frequencies of burrow establishment for Geolycosa turri- cola and G. micanopy under experimental conditions. B = burrow established, NB = no burrow established. G. turricola G. micanopy Fed Unfed Fed Unfed B NB B NB B NB B NB Vegetation 25 4 13 17 23 8 7 24 No Vegetation 20 10 9 21 22 7 12 18 Crevice — — — — 19 6 20 4 No Crevice — — — — 16 9 13 11 contingency tables. The variables were chosen because of their possible importance as factors in burrow construction based on casual and quantitative observations of laboratory and field populations of five species of Geolycosa ( G . turricola, G. micanopy, G. patellonigra, G. ornatipes, G. hubbelli and G . escambiensis). The variables were: (1) Prey/ no prey (prey) — indicating whether food was pro- vided during the experimental period. (2) Vegetation/ no vegetation (vegetation) — indicating whether small bits of grass were provided in the experiemental container. (3) Crevice/ no crevice (crevice) — indicating whether a de- pression in the burrowing surface was provided. (4) Burrow/ no burrow (burrow) — the “dependent” variable, indicating whether a burrow was constructed. Any burrow which was large enough to contain the spider was scored as an established burrow. For each experiment I tested two hypotheses concerning a three- way contingency table defined by the variables prey, burrow and one of the other two variables (Table 2). The experiments and anaylyses were performed for the two species separately for the data pertaining to the variable vegetation. No formal statistical comparison of the data for the two species was made. Analysis involving the variable crevice was carried out on G. micanopy only. The hypotheses were tested by Chi-square goodness-or-fit tests using Goodman’s (1970) maximum likelihood estimators to obtain the expected cell frequencies. 126 Psyche [Vol. 91 Sand was used as the burrowing medium in all cases. Spiderlings which were to receive food were given newly-hatched crickets at the beginning of the experimental period. A metal probe was used to make small crevices in the sand where necessary. Restricted randomization was used to assign roughly equal numbers of spiderlings to treatment groups (fed, unfed, etc.). Specimens were held in their test containers at normal room temperature and light for a period of 36 h, at which time I observed whether each had established a burrow or not. Previous observations indicated that burrows are usually established within the first 24 h after dispersal. Results The first set of hypotheses tested the independence of burrowing with the variables vegetation and crevice respectively within the levels of the variable prey (Table 2). The hypothesis that burrowing is independent of vegetation within a prey group (fed or unfed) was rejected for G. turricola but not for G. micanopy (Table 2). Both fed and unfed G. turricola showed a higher number of burrowers in the group that was provided with vegetation (86% and 43% burrowing in the fed and unfed groups respectively). For G. micanopy, the number of burrowers is approximately the same for the fed spiders with and without vegetation. A higher percentage of the unfed G. micanopy burrowed when no vegetation was present (23% and 40% for the group with and without vegetation respec- tively). More spiders burrowed in the presence of a crevice than when no crevice was available for both fed and unfed G. micanopy. The second set of hypotheses tested the independence of burrowing and prey given the level of each of the other two variables. In every case, the hypothesis of independence was rejected (Table 2). In nearly every case, the percentage of spiderlings constructing burrows was lower in the unfed group. This is true for each level of the two variables, vegetation and crevice except for the unfed G. micanopy who were provided a crevice. In that case, more spiderlings constructed burrows than did not. Discussion The results indicate that, of those variables considered, prey was most strongly associated with the establishment of a burrow. In nearly every case, groups of spiderlings which were given food had a 1984] Miller — Geolycosa 127 •O II II *- c o '■3 ^ § V c- d, ■S £ 3 O ^ Lh Z il I 6 a* 3 £ o ex £ * o 4> l_ C "fc* •- m X> O — ' 60 II a> £ ° 3 T3 ST<| u « II cx M O i| 3 rt 11 5 >- 5 o ^ <*-. II u - 2 Ui 3 CX u 0 „ , cx < ^ g u CX v O 3 § S 1 1 3 00 §•§ 'ob < U < « 5 | > .> 'ob °o DQ U U c e (L> T3 *3 C c Q Q Q Q 128 Psyche [Vol. 91 higher percentage of burrows than those that were not. Although the experimental design allows consideration of only the importance of the presence of prey, higher rates of burrow establishment in the presence of food may imply either a nutritional advantage or a response to prey availability. With respect to nutritional differences, even though burrowing behavior is innate and may be observed as early as in late pre- emergent spiderlings which have not fed (pers. observ.), the successful establishment of a burrow may be more likely if a spiderling is able to obtain food prior to dispersing. Many lycosid spiderlings disperse after only a short time on the mother (e.g., about seven days, Higashi and Rovner, 1975) and do not feed prior to dispersal (Foelix, 1982). My observations of field populations of other Geolycosa indicate that spiderlings may cling to the mother for up to three weeks and some young remain in the maternal burrow for at least one molt after leaving the mother’s abdomen as Engelhardt (1964) found in Trochosa spp. A possible advantage of this extended association with the mother is that those spiderlings that remain may receive nurishment by sharing prey captured by the mother, as in Sosippus floridanus (Brach, 1976), or by cannibalism. Hallander (1970) observed cannibalism in Pardosa pullata of the same brood, even in situations of high prey density. I held broods of G. turricola and G. patellonigra without food for six weeks and observed few instances of cannibalism. Cannibalism, therefore, is probably not a primary means of obtaining predispersal nourish- ment in Geolycosa. At present, I do not have information as to whether older juvenile Geolycosa that remain in the burrows are able to obtain food on their own. I have observed spiderlings clinging to the turret rim in a foraging position but I have never observed prey capture. The means of obtaining predispersal nourishment (if such is obtained) notwithstanding, a lack of food per se does not preclude burrow construction. Significantly more burrows were constructed in the groups which were provided with food but many unfed spiderlings (average 40.8%) successfully constructed burrows. Also, observations of lab held and starved G. patellonigra indicated that burrowing may occur well after two weeks post emergence (pers. observ.). 1984] Miller — Geolycosa 129 Riechert and Luczack (1982) recently reviewed the literature concerning the selection of microhabitat in spiders. Environmental factors such as wind (e.g., Eberhard, 1971), vegetation structure (e.g., Enders, 1975) or temperature (e.g., Riechert and Tracy, 1975) and prey characteristics such as prey availability (e.g., Kronk and Riechert, 1979; Enders, 1977; Morse, 1981) are known to influence positioning of webs and the location of foraging sites in spiders. Geolycosa burrows function in both thermoregulation (Humphreys, 1975) and prey capture (Gertsch, 1942, pers. observ.), so the placement of the burrow may be related to these functions. However, the major thermoregulatory attribute of the burrow is its depth and not its location relative to the surrounding vegetation (Humphreys, 1975). Geolycosa regulate body temperature by moving up or down the tunnel. The selection of a burrow site, therefore, is more likely to be related to prey availability and microhabitat factors relating to ease of construction or which provide some protection from predators. The results show a difference between G. turricola and G. micanopy in the relationship between vegetation availability and frequency of burrow establishment. Within a feeding state, the number of burrows established is independent of vegetation for G. micanopy but not for G. turricola. This difference reflects the turret construction habits of the two species. Geolycosa turricola nearly always constructs a conspicuous turret from whatever material is available, whereas G. micanopy shows considerable variation in turret construction and often has burrows with no turret (Wallace, 1942). The different relationship between burrowing frequency and vegetation is probably not a result of a preference for vegetation material used in the experiments, since there appears to be no specificity for turret material in Geolycosa (Wallace, 1942). Nearly all Geolycosa observed in the lab readily used artificially constructed burrows. Field observations of dispersing G. turricola (Miller and Miller, in prep.) indicate that over one half of burrows constructed by dispersing G. turricola spiderlings were built within a surface crack or depression. The results presented here also indicate a preference for burrowing when a surface irregularity is present. Surface cracks and crevices could provide protection from pred- ators and thermoregulatory advantage during the initial phase of 130 Psyche [Vol. 91 burrow construction. They may also give the spider a foraging advantage by providing an ambush location. Evidence of the importance of surface cracks is given by the unfed G. micanopy which were provided with cracks. That group constructed nearly as many burrows as the fed group. Summary The relationship between the establishment of a burrow and presence of prey, availability of vegetation and the presence of a crevice in the burrowing surface was investigated in newly- dispersing Geolycosa turricola (Treat) and G. micanopy Wallace. The establishment of a burrow by G. turricola was dependent on the presence of vegetation within a feeding group. Establishment of burrows by G. micanopy was dependent on the presence of a crevice (not tested for G. turricola). Burrow establishment was found to be dependent on prey availability for any level of the other variables. The dependence on vegetation for burrow establishment in G. turricola is attributed to a greater tendency to build turrets in that species. The higher burrowing frequency in the groups that received food during the experiment is thought to be related to nutrition and/or prey availability. The tendency to build new burrows in crevices is known from field studies and may be related to advantages of protection from predators, thermoregulation, or foraging position imparted by the crevice. Acknowledgments I appreciate the thoughtful comments of J. Rovner, G. Baker, M. LaSalle, R. Altig, M. Morris, P. Ramey-Miller and several anonymous reviewers on various drafts of this paper. Literature Cited Brach, v. 1976. Subsocial behavior in the funnel-web wolf spider Sosippus floridanus (Araneae:Lycosidae). Fla. Ent. 59(3):225-229. Eberhard, W. G. 1971. The ecology of the web of Uloborus diversus (Araneae: Uloboridae). Oceologia 6(4):328-342. Enders, F. 1975. The influence of hunting manner on prey size, particularly in spiders with long attack distances (Araneidae, Linyphiidae, and Salticidae). Amer. Natur. 901(970):737-763. 1984] Miller — Geolycosa 131 1977. Web-site selection by orbweb spiders, particularly Argiope aurantia Lucus. Anim. Behav. 15:694-712. Englehardt, W. 1964. Die mitteleuropaischen arten der gattung Trochosa C. L. Koch 1848 (Araneae, Lycosidae). Morphologie, chemotaxonomie, biologie, auto- kologie. Z. Morph. Okol. Tiere 54:219-393. Foelix, R. F. 1982. Biology of Spiders. Harvard University Press. Cambridge, Massachu- setts. 306 p. Gertsch, W. J. 1949. American Spiders. D. van Nostrand Co., Inc. New York. 285 p. Goodman, L. A. 1970. The multivariate analysis of qualitative data: Interactions among multiple classifications. J. Amer. Stat. Assoc. 65:226-256. Hallander, H. 1970. Prey, cannibalism and microhabitat selection in the wolf spider Pardosa chelata O. F. Muller and P. pullata Clerck. Oikos 21:337-340. Humphreys, W. F. 1975. The influence of burrowing and thermoregulatory behavior on the water relations of Geolycosa godeffroyi (Araneae: Lycosidae) an Australian wolf spider. Oecologia 21(4):29 1 —3 1 1 . Higashi, G. A. and J. S. Rovner. 1975. Post-emergent behaviour of juvenile lycosid spiders. Bull. Brit. Arch. Soc. 3(5): 1 13—1 19. Krebs, C. J. 1978. Ecology: The experimental analysis of distribution and abundance, second edition. Harper and Row, Publishers. New York. 678 p. Kronk, A. W. and S. E. Riechert. 1979. Parameters affecting the habitat choice of Lycosa santrita Chamberlin and Ivie. J. Arachnol. 7:155-166. McCrone, J. D. 1965. Geographical variation in the seasonal distribution of Geolycosa patellonigra (Araneae, Lycosidae). Am. Mid. Nat. 73(1): 166- 169. McQueen, D. J. 1978. Field studies of growth, reproduction, and mortality in the burrowing wolf spider Geolycosa domifex (Hancock). Can. J. Zool. 56:2037-2049. Morse, D. H. 1981. Prey capture by the crab spider Misumena vatia (Clerck) (Thomisidae) on three common native flowers. Amer. Midi. Natur. 105(2):358— 367. Riechert, S. E. 1976. Web-site selection in a desert spider Agelenopsis aperta (Gertsch). Oikos 27:311-315. Riechert, S. E. and C. R. Tracy. 1975. Thermal balance and prey availability: bases for a model relating web- site characteristics to spider reproductive success. Ecology 56:265-284. Riechert, S. E. and J. Luczak. 1982. Spider foraging: Behavioral responses to prey, in P. N. Witt and J. S. 132 Psyche [Vol. 91 Rovner. eds.. Spider Communication: Mechanisms and Ecological Significance. Princeton University Press. Princeton, New Jersey. 236 pp. Savory, T. H. 1930. Environmental differences of spiders of the genus Zilla. J. Ecol., 18:384-385. Waldorf, E. S. 1976. Spider size, microhabitat selection, and use of food. Am. Mid. Nat. 96(l):76-87. Wallace, H. K. 1942. A revision of the burrowing spider of the genus Geolycosa (Araneae; Lycosidae). Am. Mid. Nat. 27( 1 ): 1 —6 1 . RECENTLY RECOGNIZED TYPES OF SOME HOMOPTERA DESCRIBED BY DR. ASA FITCH' By Jeffrey K. Barnes Biolological Survey New York State Museum The State Education Department Albany, NY 12230 Dr. Asa Fitch (1809-1879) proposed 165 new species and subspecies names in the Homoptera. They are taxonomically distributed as follows: Cicadidae (2), Membracidae (21), Cicadelli- dae (34), Cercopidae (9), Delphacidae (2), Derbidae (7), Cixiidae (2), Achilidae (2), Issidae (2), Psyllidae (9), Aleyrodidae (1), Aphididae (52), Adelgidae (2), Phylloxeridae (4), Diaspididae (5), and Coccidae (11). He proposed the names in print from 1851 to 1872, mainly in his Homoptera catalogue (1851) and the Trans- actions of the New York State Agricultural Society. Dr. Fitch never designated types for his new species and subspecies. However, he did label nearly all of his specimens with individual numbers. He recorded these numbers consecutively in four individual registers, along with collecting dates, localities, and other pertinent information. Label numbers in black ink written on white paper are recorded in one register of specimens from New York State. Label numbers in black ink crossed with one or two red lines and written on white paper are recorded in a second register of specimens from New York State. One red line indicates a number less than 10,000, and two red lines indicate a number to which 10,000 should be added. Label numbers in red ink written on white paper are recorded in a third register that lists specimens from elsewhere in North America. These three registers are in the possession of the New York State Museum. The fourth register, owned by the Museum of Science, successor to the Boston Society of Natural History, lists specimens from elsewhere in the world. The corresponding labels are written in black ink on colored papers. Dr. Fitch also kept individual sheets of notes on every species of which 'Published as New York State Museum Journal Series No. 426. Manuscript received by the editor February 20, 1984. 133 134 Psyche [Vol. 91 he was aware. If he had specimens of any one species, he recorded the specimen label numbers along with collection data on the appropriate manuscript sheet. Most of these notes are in the New York State Museum. Many of the Homoptera notes are in the Smithsonian Institution Archives and the files of the United States National Museum aphid collection. According to the International Code of Zoological Nomen- clature, the type series of a species or subspecies consists of all the specimens on which its author bases the species (Article 72b). The closest available approximation to a type series for a species or subspecies authored by Dr. Fitch consists of those specimens that can be proven, through a perusal of his registers and notes, to have been in his collection before or during the year in which the relevant species or subspecies name was first made available. In the introduction to his Homoptera catalogue, dated February 22, 1851, Dr. Fitch acknowledged the loan of all of Dr. Thaddeus William Harris’s collection of Homoptera. In a letter to Dr. Harris, Dr. Fitch stated that he was forwarding copies of his Homoptera catalogue. He also stated that it would be a pleasure for him to add specimens from his collection to Dr. Harris’s collection.2 That letter is dated March 1-4, 1851. The T. W. Harris insect collection at the Museum of Comparative Zoology, Harvard University, is segregated from the main collec- tion. In it, several of Dr. Fitch’s specimens are readily recognized by the label style and handwriting (Fig. 1). Some of these are specimens of species that Fitch described in his Homoptera catalogue. Those that were in Dr. Fitch’s possession before February 28, 1851 (the earliest publication date for the Homoptera catalogue that can be demonstrated by evidence) can be considered types, but they have never been recognized as such. In most cases, these specimens are part of a syntype series, and other specimens can be found in the New York State Museum (McCabe and Johnson 1980) or the United States National Museum. In these cases, lectotypes should be designated if that has not already been done. Other specimens in the Harris collection are the only known extant specimens from the Fitch collection that Dr. Fitch considered to be representatives of certain species that he authored. 2 Fitch to Harris, 1-4 March, 1851; Museum of Comparative Zoology, Harvard University. 1984] Barnes — Types of Homoptera 135 o If# 2 f/f? f 7? j?f#s £?$/ jyfTi. b y? 2ffs iff* Jffy -2. Fig. 1. Dr. Asa Fitch’s handwriting and style of insect labels, a, uncut strip of labels; b, close-up view of a section of a; c, New York insect label 14,059, black ink on white paper; d, New York insect label *9909, black ink on white paper, crossed with one red line. Labels c and d same scale. The following list contains brief descriptions of specimens from Dr. Fitch’s collection that can now be found in the Harris collection. Generic and specific names are spelled exactly as they were in the original publications. Only specimens of species authored by Dr. Fitch are listed. The label numbers that are preceeded by one or two asterisks correspond with specimen labels that are written in black ink on white paper and crossed with one or two red lines, respectively. A summary of collecting data, derived from Dr. Fitch’s registers and notes, accompanies each description of a specimen. 136 Psyche [Vol. 91 Membracidae Uroxiphus caryae Fitch, 1851:52. *3992, Greenwich, NY, 24.viii.1846. *3997, Greenwich, NY, 24.viii.1846. Tragopa dorsalis Fitch, 1851:52. **134, Salem, NY, 25.vii.1851. **137, Salem, NY, 25.vii.1851. Cyrtoisa fenestrata Fitch, 1851:49. *6927, Long Island, NY, 1847. ClCADELLIDAE ldiocerus alternatus Fitch, 1851:59. *8665, Salem, NY, 7.V.1851. *8828, Salem, NY, lO.v.1851. *9075, Salem, NY, 19.V.1851. Helochara communis Fitch, 1851:56. *8761, Salem, NY, lO.v.1851 Amblycephalus curtisii Fitch, 1851:61. 12,702, Salem, NY, 22.xi.1850. Gypona flavilineata Fitch, 1851:57. *6304, Salem, NY, 10.ix.1847. Jassus fulvidorsum Fitch, 1851:62. *6178, Salem, NY, 2 1 . viii. 1 847. Amblycephalus melsheimerii Fitch, 1851:61. *4911, Salem, NY, 5.vii.l847. Athysanus minor Fitch, 1851:60. *4869, Salem, NY, 5.vii.l847. Athysanus nigrinasi Fitch, 1851:61. *2983, Jackson, NY, 16.vi.1846. Aulacizes noveboracensis Fitch, 1851:56. *9936, Salem, NY, 12.vii.1851. ldiocerus pallidus Fitch, 1851:59. *6059, Salem, NY, 19.viii.1847. *9834, Salem, NY, 5.vii.l851. Empoa querci Fitch, 1851:63. 13,299, Stillwater, NY, vii.1848. Amblycephalus sayii Fitch, 1851:61. *9234, Salem, NY, 30.V.1851. 1984] Barnes — Types of Homoptera 137 Bythoscopus strobi Fitch, 1851:58. *9397, Salem, NY, 5.vi.l851. Erythroneura tricincta Fitch, 1851:63. *6402, Salem, NY, 10.ix.1847. Bythoscopus unicolor Fitch, 1851:58. 2021, Tullehassie, AR [Tullahasee, OK], viii.1851. *4909, Salem, NY, 5.vii.l847. Athysanus variabilis Fitch, 1851:60 *9822, Salem, NY, 5.vii.l851. Pediopsis viridis Fitch, 1851:59. *9836, Salem, NY, 5.vii.l851. **153, Salem, NY, 25.vii.1851. Erythroneura vulnerata Fitch, 1851:62. *6467, Salem, NY, 10.ix.1847. *6621, Greenwich, NY, 20.ix.1847. *6629, Greenwich, NY, 20.ix.1847. Cercopidae Clastoptera testacea Fitch, 1851:53. *5322, Salem, NY, 23.vii.1847. Delphacidae Delphax arvensis, 1851:46. 12,358, Salem, NY, 3.vi.l847. Delphax dorsalis Fitch, 1851:46. *8500, Salem, NY, 26.iv.1851. *8578, Salem, NY, 29.iv.1851. ClXIIDAE Cixius pini Fitch, 1851:45. 474, Winhall, VT, 17.vi.1847. ISSIDAE Bruchomorpha dorsata Fitch, 1857:396. 1555, Tullehassie, AR [Tullahassee, OK], l.vi.1851. Naso robertsonii Fitch, 1857:396. 1776, Tullehassie, AR [Tullahassee, OK], vii.1851. 138 Psyche [Vol. 91 PSYLLIDAE Psylla carpini Fitch, 1851:64. *9224, Salem, NY, 30.V.1851. Aphididae Aphis crataegifoliae Fitch, 1851:66. *9287, Salem, NY, 2.vi.l851. Aphis populifoliae Fitch, 1851:66. *9294, Salem, NY, 5.vi.l851. *9298, Salem, NY, 5.vi.l851. Eriosoma pyri Fitch, 1851:68. *4053, Salem, NY, 10.xi.1846. *4054, Salem, NY, 10.xi.1846. Aphis rudbeckiae Fitch, 1851:66. 11,715, Salem, NY, 13.vii.1846. 11,717, Salem, NY, 13.vii.1846. Dr. Fitch described six new genera, 85 new species, and five new subspecies in his Homoptera catalogue. The New York State Museum’s collection still contains type specimens of 60 of the species and subspecies. The remainder have been considered destroyed (Lintner 1893, McCabe and Johnson 1980). The speci- mens that Dr. Fitch listed in his catalogue were numbered 609-874, and he tagged the specimens with numbered labels printed expressly for that purpose (Funkhouser 1915). These numbers bear no relationship to the numbers in Dr. Fitch’s manuscript species notes or specimen registers. In the United States National Museum there are six specimens labelled in Dr. Fitch’s handwriting with three digit numbers that correspond with those in his Homoptera catalogue. They are two specimens of Psyllidae, Livia femoralis Fitch (838) and L. vernalis Fitch (836), and four specimens of Aphididae, Aphis betulaecolens Fitch (848), A. cerasicolens Fitch (841), A. berberidis Fitch (842), and A. sambucif oliae Fitch (850). Specimen 842 is also labelled Aphis pinicolens Fitch, but not in Fitch’s handwriting. A seventh specimen in the USNM — a specimen of Aphis cornifoliae Fitch — is labelled “846 Fitch,” “Type,” and “Female,” corre- sponding with the information given in Fitch’s catalogue, but these labels are not in Fitch’s handwriting. Two more specimens of Aphididae, Lachnus abietis Fitch (854) and L. alnifoliae Fitch (857), 1984] Barnes — Types of Homoptera 139 are described in Smith and Parron’s list of North American Aphididae (1978) as “nearly destroyed, in USNM,” but I have been unable to locate them. Literature Cited Fitch, A. 1851. Catalogue with references and descriptions of the insects collected and arranged for the State Cabinet of Natural History. Pages 43-69 in Fourth annual report of the Regents of the University, on the condition of the State Cabinet of Natural History, and the historical and antiquarian collection, annexed thereto. Made to the Senate, January 14, 1851. Albany. 146 pp. [1857]. Third report on the noxious and other insects of the State of New York. Transactions of the New York State Agricultural Society 16(1856): 315-490. Funkhouser, W. D. 1915. Types of Fitch’s species of Membracidae. Bulletin of the Brooklyn Entomological Society 10: 45-50. Lintner, J. A. 1893. Catalogue of the known Homoptera of the State of New York in 1851. Annual Report of the New York State Museum 46: 381. McCabe, T. L., and L. M. Johnson. 1980. Catalogue of the types in the New York State Museum insect collection. New York State Museum Bulletin 434: 1-38. Smith, C. F., and C. S. Parron. 1978. An annotated list of Aphididae (Homoptera) of North America. North Carolina Agricultural Experiment Station Technical Bulletin 255: 1-428. PRODUCTION AND USE OF SECRETIONS PASSED BY MALES AT COPULATION IN PIERIS PROTODICE (LEPIDOPTERA, PIERIDAE)* By Ronald L. Rutowski1 Department of Zoology Arizona State University Tempe, AZ 85287 Introduction During copulation in many species of insects the male passes to the female sizeable quantities of accessory gland secretions with the sperm (Thornhill and Alcock, 1983). Recently it has been shown in the Lepidoptera that these secretions may represent a nutrient investment by the male that is used by the female in the production of eggs and in somatic maintenance (Boggs and Gilbert, 1979; Boggs, 1981; Boggs and Watt, 1981; Goss, 1977; Greenfield, 1982). Typically these secretions are contained within a spermatophore with the sperm or they may be passed as loose secretion. The perspective that these secretions may constitute a nutrient invest- ment by males has some important implications for how male and female lepidopterans should behave (Marshall, 1982; Rutowski, 1982; see also, Gwynne, 1982). In spite of recent interest in these secretions there are few studies of the patterns of their production and use within and between species of Lepidoptera (e.g., Boggs, 1981; Greenfield, 1982; Rutow- ski et al., 1983). This paper reports data on patterns of variation in the quantity of material passed to females during copulation by males of the checkered white butterfly ( Pieris protodice Boisduval and LeConte) and the disappearance of these secretions from the female’s reproductive tract. The secretions are received by the female in the bursa copulatrix which in this butterfly species is composed of two sacs, the corpus bursa and the appendix bursa. These are connected in series with the appendix bursa being furthest from the copulatory opening. During copulation the appendix bursa is filled with a whitish fluid and the corpus bursa is filled with * Manuscript received by the editor March 5, 1984. 141 142 Psyche [Vol. 91 a spermatophore that is formed within the bursa during copulation and contains sperm and secretions. Methods and Materials All observations were made at the Arizona State University Farm Laboratory from March to July in 1983. Here the butterflies use cultivated alfalfa as a nectar source and feral and cultivated crucifers as oviposition sites and larval foodplants.All hand-reared adults were grown from eggs collected in the laboratory or from young larvae collected in the field. In the laboratory larvae were reared either on broccoli ( Brassica spp.) leaves or on cuttings of Sisymbrium irio L. in plastic shoe boxes under variable and uncontrolled conditions of light and humidity. The temperature in the laboratory was about 24° C. Production of secretions during copulation To examine the patterns of production of secretions by males, virgin females were released next to free-flying males and permitted to copulate with them. If the copulation lasted 60 min or less, both members of the pair were freeze-killed after the pair separated. Longer copulations were thought to indicate dysfunction or recent mating by the male. Males in the mated pairs were assessed with respect to wing wear (age), forewing length, and body weight (to the nearest 0.1 mg) (for techniques, see Rutowski et al., 1983). Females were assessed with respect to forewing length and the volume and wet mass of material received from the male during copulation (for technique see Rutowski et al. 1983). In some cases males were weighed and measured without killing so that they could be held overnight (without food and water) and returned to the field the following day to observe their performance in second matings. Male precopulatory body mass was estimated by adding the mass of the secretions imparted to his postcopulatory mass. To examine the rate of use of the material received during copulation virgin females were mated to free-flying males as above and then held in cylindrical screen cages (9 cm diameter, 12 cm deep) in the laboratory under flourescent illumination. At various times after mating females were freeze killed and dissected to examine the material remaining in the bursa copulatrix. Sprigs of S. irio for oviposition and sugar water for food were available in these cages. 1984] Rut ow ski — Pier is protodice 143 All summary statistics are given as mean ± standard deviation. The 0.05 level was used in all evaluations of statistical significance. Results Production of secretions by males Various aspects of the observed copulations and the individuals that participated in them are summarized in Table 1. Again, all these copulations were less than 60 min in duration. All males were in fresh or only slightly worn condition. Several significant positive correlations reflect concordance in the measuring techniques. Male forewing length was significantly positively correlated with the estimate of male precopulatory body mass (Fig. 1A; r=0.71, n=26, p< 10 s) and a similar relationship was found between the volume and the mass of male-imparted secretions (Fig. IB; r= 0.879, n=28, p<109). Qualitative microscopic observations were made on squash mounts of material removed from the spermatophore and the ap- pendix bursa found in 3 females. These observations suggest that the sperm packet is (1) discrete from the non-sperm matieral, (2) last to be deposited in the bursa copulatrix (closest to the bursal entrance), and (3) typically consititues 10% or less of the volume of material passed by the male. On average males passed almost 8% of their precopulatory body mass at copulation. However, the quantity of material passed measured by volume or mass was not significantly correlated with either male forewing length (vs. volume: r=-0.054, n=28, p=0.39; vs. mass: r=— 0.108, n=28, p=0.29) or male precopulatory body mass (vs. volume: r= -0.095, n=26, p=0.32; vs. mass: r=0.0066, n=26, p=0.48). The percent of male precopulatory body mass passed at copulation was significantly negatively correlated with both measures of male size (Fig. 2; forewing length: r=-0.625, n=26, p=0.0003; precopulatory body mass: r=-0.654, n=26, p= 0.0001). These results suggest that all males regardless of size pass a typical amount of material. Small males must then obviously pass a larger proportion of their body mass to achieve this typical quantity. Copulation duration was not significantly correlated with the volume of material passed (r=-0.243, n=26, p=0.115) but was significantly negatively correlated with the mass of secretions passed by males (Fig. 3; r=-0.34, n=26, p=0.045). There may be a 144 Psyche [Vol. 91 Table 1. Summary of 28 copulations that lasted 60 min or less. Variable (units) Mean Standard deviation Sample size Range Male forewing length (mm) 23.9 1.1 28 21-26 Estimate of male precopulatory body mass (mg) 52.9 10.9 26 29.1-70 Female forewing length (mm) 23.3 1.49 28 19-26 Copulation dura- tion (min) 31.7 9.23 26 22-60 Volume of male-imparted secretions (/u) 3.44 1.08 28 1.36-5.46 Mass of male-imparted secretions (mg) 3.89 1.15 28 1.78-5.9 % of male body mass passed 7.88 2.59 26 2.7-12.7 tendency in copulations lasting an hour or less for females to receive smaller quantities of material than in long copulations. Female forewing length was not significantly correlated with male forewing length (r= -0.0404, n=28, p=0.42) which suggests size assortative mating does not occur in this species. Female forewing length was not significantly correlated with the volume of secretions received (r=-0.25, n=28, p=0.10) or the mass of secretions received (r=0.228, n=26, p=0.122). Female size apparently does not in- fluence the quantity of material passed. Ten males were mated and then successfully remated 24 hours later. Table 2 summarizes a comparison of first and second matings by these males. Statistical comparisons reveal that significantly less material was passed in the second matings (t = 4.32, 9 df, p<0.002) and males passed a significantly smaller proportion of their body mass (t=3.3, 9 df, p<0.01). Although not all copulation durations were precisely measured, those that were displayed no difference between the durations of first and second matings separated by 24 hrs (t = 0. 1 07, 15 df, p=0.46). Rate of use of male secretions by female Fig. 4 shows the mass of material remaining in the females’s reproductive tract at various times after mating. During the first day or two after copulation little material has left the bursa copulatrix but then the decline is rapid until day 5 at which point it seems to level off. The correlation between days since copulation and ma- terial left in the bursa copulatrix was negative and highly significant (r=— 0.751, n = 46, p<10"8). 1984] Rutowski — Pieris protodice 145 (irf) aiuniOA • • • •% -LO _C CM O) CM c d) < • • i ■ — i > — i — -» — i o o o o oo co cm (6uu) ssduu Apog £ g> _c\j $ CM d) £ Figure 1. Concordance in measures of (A) male size and (B) quantity of material passed at copulation. 146 Psyche • • • • I1 I 1 I"1 1 1 T "> — l — r •• • •• •• %• •• 9 / • • • o 00 — U) E o oE ■D O GO o C\J 8? ™x: D) CM £ CO CM CM CM U) c I ' I 1 I • I * I * 1 • I ■“ I ^TCMOOOCDmT^O passDd ssduu Apoq °/Q £ O 2 — 61 a; § O)4' §3- I 2‘ 0 1- • • • • rxi/ 0J • 1 ' 1 1 1 1 1 r~ 01 234567 >7 Days after copulation Figure 4. The quantity of material remaining in the female’s bursa copulatrix as a function of the time since copulation. Line shows change in mean values for days 0 through 7. and second matings separated by 24 hrs (Table 2). Several compari- sons are valuable here. First, there was no significant difference between the amount of material passed in first copulations in 1979 and 1983 (t = 0.614, 26 df, 0=0.27). In addition, males passed significantly more material 24 hrs after the first mating than they did 10 min after the first mating (t=2.95, 17 df, p= 0.004). These comparisons point out the consistency in the observations between the two years and more importantly that while copulation durations indicate that males have substantially recovered in 24 hr their supply is still somewhat depleted. However, this depletion may be a result of housing males between matings without food and water. In fact most of these males flew immediately to nectar sources upon release in the field. There is much to be learned about the performance of males in third matings and their performance in second matings when they have access to nectar. Use of secretions by females The rate at which females deplete the secretions they have received from males is greatest between the second and fifth days 1984] Rut ow ski — Pier is protodice 151 after mating. This suggests that a female’s supply of secretions but not sperm is effectively depleted beginning at about six days after mating. Suzuki (1979) examined the phenomenon of multiple mating in Pier is rapae L., a close relative of P. protodice, and found that most females mate for the second time about 6 to 8 days after being released as a virgin in the field. These data support Suzuki’s hypothesis that Pieris females may multiply mate to gain nutrients in that they appear to be doing so at a time when their supply of secretions from previous matings is largely depleted. This hypothe- sis is also confirmed by the data from the dissections of field- collected females in which at least one of the spermatophores carried by twice-mated females was often small and greatly depleted. Summary This study examines the production and use of secretions passed by a male during copulation to a female in the checkered white butterfly ( Pieris protodice). Males pass about 7 to 8 percent of their body weight at copulation in the form of secretions deposited in the female’s reproductive tract. No correlation between the quantity of material passed and male size was found. However, it was found that the proportion of male body mass passed was inversely correlated with body mass suggesting that all males, regardless of size pass a typical quantity of material. A mating depletes the material a male has for subsequent mating but only temporarily. Males were found to have mostly recovered their potency in 24 hrs, even without food or water between matings. Females have mostly depleted the material received from males in about 5 to 7 days. In the field females carrying multiple sperm- atophores always had one that was greatly depleted indicating that they remate only when material from a previous mating was depleted. The results are compared to information on the use and production of male-imparted secretions and mating behavior in other butterflies. Acknowledgements I thank John Schaefer and Scott Snead for their assistance in the field and laboratory. Financial support was provided by National Science Foundation Grants BNS 80-14120 and BNS 83-00317. 152 Psyche [Vol. 91 References Boggs, C. 1981. Boggs, C. 1979. Boggs, C. 1981. Goss, G. L. Selection pressures affecting male nutrient investment at mating in heliconiine butterflies. Evolution 35: 931-940. L. and L. E. Gilbert. Male contribution to egg production in butterflies: evidence for transfer of nutrients at mating. Science 206: 83-84. L. AND W. B. Watt. Population structure of pierid butterflies. IV. Genetic and physiological investment in offspring by male Colias. Oecologia 50: 320-324. 1977. The interaction between moths and pyrrolizidine alkaloid-containing plants including nutrient transfer via the spermatophore in Lymire edwardsii (Ctenuchidae). Ph.D. Thesis, University of Miami. Gwynne, D. T. 1982a. Male nutritional investment and the evolution of sexual differences in Tettigoniidae and other Orthoptera. Pages 337-366 in D. T. Gwynne and G. K. Morris, ed., Orthopteran Mating Systems: Sexual Compe- tition in a Diverse Group of Insects. Westview Press, Boulder. 1982b. Mate selection by female katydids (Orthoptera: Tettigoniidae, Cono- cephalus nigropleurum). Anim. Behav. 30: 734-738. Greenfield, M. D. 1982. The question of paternal investment in Lepidoptera: male-contributed proteins in Plodia inter punctella. Int. J. Invert. Reprod. 5: 323-330. Marshall, L. D. 1982. Male nutrient investment in the Lepidoptera: what should males invest? Amer. Nat. 120: 273-279. Rutowski, R. L. 1979a. Courtship behavior of the checkered white, Pieris protodice (Pieridae). J. Lep. Soc. 33: 42-49. 1979b. The butterfly as an honest salesman. Anim. Behav. 27: 1269-1270. 1982. Mate choice and lepidopteran mating behavior. Florida Ent. 65: 72-82. Rutowski, R. L., M. Newton and J. Schaefer. 1983. Interspecific variation in the size of the nutrient investment made by male butterflies during copulation. Evolution 37: 708-713. Sugawara, T. 1979. Stretch reception in the bursa copulatrix of the butterfly, Pieris rapae crucivora, and its role in behaviour. J. Comp. Physiol. 130: 191-199. Suzuki, Y. 1978. Adult longevity and reproductive potential of the small cabbage white, Pieris rapae crucivora Boisduval (Lepidoptera: Pieridae). Appl. Ent. Zool. 13: 312-313. Thornhill, R. and J. Alcock. 1983. The Evolution of Insect Mating Systems. Harvard University Press, Cambridge. FEMALE MONOGAMY AND MALE COMPETITION IN P HO TIN US COLLUSTRANS (COLEOPTERA: LAMPYRIDAE)* By Steven R. Wing Department of Entomology and Nematology University of Florida Gainesville, Florida 32611 Introduction Matings by Photinus collustrans females can easily be kept track of in the field. These brachypterous females live in burrows and remain near them. About 20 minutes after sunset males start to fly and search for females, which take positions on the soil surface or on vegetation (Lloyd 1966). Females flash in response to the signals of flying males, which locate them by their responses. Each night sexual activity is restricted to a period about 18 minutes long (Lloyd 1966; also see T. Walker 1983 for a discussion of such ‘sprees’). P. collustrans females live about 10 days after their first appearance (Wing 1982); by observing a female for about 20 minutes per night for 10 nights, every sexual activity of her life can be recorded. There is no evidence that females mate under circumstances other than those mentioned above. This field study shows that only a very small proportion of collustrans females mate more than once. Yet the potential for female multiple mating apparently is the basis for the evolution of a complex of tactics used by competing males. Methods and Materials Field studies of collustrans were conducted in Alachua County, Florida. The site was a grassy area under scattered oaks, pines, and shrubs. The grass was mowed periodically. Two streetlights illuminated parts of the site. Females were located by their flash and/or glow responses to penlight simulations of male mating signals (see Lloyd 1966). The location of each female was marked by placing a numbered flag about 15 cm to the north of her. Flags were 10 X 40 cm strips of * Manuscript received by the editor January 22, 1984. 153 154 Psyche [Vol. 91 plastic held in place with nails. Females use the same burrow throughout their adult lives1 (Wing 1982), and numbered flags placed at the burrow location were sufficient to identify each individual. Copulations were timed with a stopwatch, and some were observed with a magnifying glass. An 18 X 20 m area was searched for females nightly starting before male flights began and ending after they had ended for the night. The location of each female was marked and I inspected her position at approximately 1 min intervals. Each time a female’s position was visited on a given night, she was presented penlight simulations of the male signal. Because females do not respond following a successful mating, but pause and then re-enter their burrow (Wing 1982), a female answer to my signal indicated that she was not yet mated. If she failed to respond, I determined whether she was 1) still out but not responsive; 2) mating; 3) entering her burrow; or 4) gone. All female locations were checked until the adult season was over. Because the same area was searched nightly throughout the season, when a new female appeared she was almost certainly a virgin making her first appearance. Complete sexual histories of 108 collustrans females were recorded. Results Of the 108 females whose complete sexual histories were determined, 104 mated only once. The general sexual pattern was as follows. The female appeared by her burrow nightly until she attracted a male (x = 2 nights). The male, having located the female by her continued responses to his signals, landed nearby and walked to her. Upon making physical contact, the male climbed upon the female, and copulated with her in the male-above position (Fig. 1). Copulation lasted about 1 min (details below). The male broke the connection, dismounted, and flew away leaving the female outside her burrow. Following copulation the female paused for seconds or minutes, and did not flash responses to signals of passing males. She then entered her burrow. 'During this study 91 females were individually marked with Tech Pen Ink dispensed from Hamilton’s paint pots (T. Walker and Wineriter 1981). Females appeared for up to 10 consecutive nights. Every appearance by each marked female occurred at her original position. 1984] Wing — Photinus collustrans 155 Four females mated more than once. One of these females was dug from her burrow by a male, one was mated by a “sneaky” male, and two made themselves available to males by their own behavior. The two repeated matings due to female behavior occurred when females mated and entered their burrows, but on subsequent nights left their burrows, responded to male signals, and mated again. Only three of 108 females appeared on nights subsequent to the first mating. Two mated again; one remated once, the other twice. When more than one male landed at a responding female, the first male to reach her mounted and began copulation. The rival male attempted to mount the female (sometimes backwards), and to break the pair apart (Fig. 2). As a result, the copulating male moved or was pushed off the female and copulations proceeded in the tail- to-tail position, and variations thereof. (In the tail-to-tail position the male and female face in opposite directions while maintaining genitalic connection. Due to disturbance by rival males, pairs were sometimes moved into odd positions, even with the female on the copulating male’s dorsum.) Copulations were significantly longer when rival males were present. Mean duration for single male copulations was 57 sec (n=23, range 30-185 sec) compared to 842 sec (n=5, range 339-1410 sec) Mann-Whitney (U=l 15) P < .0005 when rivals were present. In these cases, copulating males maintained the genital connection until after the females had entered their burrow (Fig. 3). Females entered head first, dragging the coupled males backwards down the burrow. In one case, only the head and thorax of the male remained outside the burrow when the genital connection was broken. After disengaging, males climbed out and flew away. After the mated male departed, in four of seven cases the rival male tried to remove the female from her burrow. Rival males located the burrow opening by antennating the soil. Rivals dug at the burrow (Fig. 4), sometimes completely entering it. On one occasion the male succeeded in removing the female from her burrow and mated with her (this accounted for the third multiple mating) (Fig. 5). Unsuccessful males dug for as long as 35 min before leaving. The fourth repeated mating resulted from another behavior of rival males and was observed once during this study and once since then. The rival “sneaky” male was non-aggressive, and made only occasional contact with the copulating pair. The rival gently antennated the pair and then walked away, returned and antennated 156 Psyche [Vol. 91 Fig. 1. Photinus collustrans male (above) mounting female. Note burrow opening in background. The female is about 1 1 mm long. Fig. 2. Photinus collustrans trio. Male on left is copulating. Rival has mounted female. 1984] Wing — Photinus collustrans 157 Fig. 3. Rival male (left) remains mounted as female enters burrow. Male on right is copulating. Fig. 4. Rival male digs at female’s burrow. 158 Psyche [Vol. 91 again. The copulating male stayed in the male-above position, and copulation was not prolonged. After copulation, the male dis- mounted and flew away, leaving the female outside her burrow. The rival male then located the female and mated with her. Discussion Females are difficult for males to locate (Lloyd 1979). With a period of only about 18 min nightly to operate in, the usual male strategy after finding a female is to mate and, within a minute or so, return to the air searching for another female. Females pause after mating, but do not answer the flashes of passing males. They then enter the burrow. Less than 3% of the females that mated ever made themselves available to males again (also see Wiklund 1982), and those that did may have had some fault in the mechanism that indicates whether sufficient sperm was acquired (also see W. Walker 1980). During the pause before re-entering the burrow females are susceptible to another mating if found. Even after re-entering the burrow a female may be dug up and remated. Generally, then, if a male can gain physical access to a female, he can mate with her. The fact that an accessible female could be mated if found has led to prolonged copulation when a rival male is present (Parker 1970). Copulating males make the female physically unavailable by occupying her until she has returned to her burrow (see Sivinski 1983). Rival males try to break the coupled pair apart and attempt to gain access to the female by digging her from the burrow. The “sneaky” rival avoids triggering mate-guarding by the coupled male, and thereby gains access to the female after her first mate leaves. The frequency of male encounters with rivals might vary with male density, but the overall proportion of females that mate repeatedly is probably rarely if ever much greater than the four per hundred found in this study.2 The complex of male strategies and counter-strategies shown here reflects how important the potential for female multiple mating can be, even when only a small proportion of females actually mate more than once. 2 Based on 71 matings observed when male density was at a 3-year peak (Wing unpublished data). It should also be noted that mated females are sometimes flooded from their burrows, and may remate under these circumstances (Wing 1982 and unpublished). 1984] Wing — Photinus collustrans 159 Fig. 5. Rival male pulls female from her burrow. Summary Four of 108 Photinus collustrans females mated more than once. One was dug from her burrow by a male, one was mated by a “sneaky” male, and two made themselves available to males after mating. Acknowledgments The photograph for Fig. 2 was made by T. G. Forrest. J. E. Lloyd introduced me to fireflies and reviewed the manuscript several times. Reviews by T. G. Forrest and J. Sivinski also improved the paper. Special thanks for comments by members of the Editorial Board at Psyche. Barbara Hollien typed the manuscript. S. A. Wineriter helped mount the photographs. Some of the work was funded by NSF grant no. DEB-7821744. Florida Agricultural Experiment Station Journal Series No. 4934. Literature Cited Lloyd, J. E. 1966. Studies on the flash communication system in Photinus fireflies. Univ. Mich. Misc. Publ. No. 130. 160 Psyche [Vol. 91 1979. Sexual selection in luminescent beetles. Pages 293-342 in M. S. Blum and N. A. Blum (eds.): Sexual selection and reproductive competition in insects. Academic Press, New York. Parker, G. A. 1970. Sperm competition and its evolutionary consequences in the insects. Biol. Rev. 45:525-567. SlVINSKI, J. 1983. Predation and sperm competition in the evolution of coupling durations, particularly in the stick insect Diapheromera veliei. Pages 147-162 in D. T. Gwynne and G. K. Morris (eds.): Orthopteran mating systems: sexual competition in a diverse group of insects. Westview Press, Boulder. Walker, T. J. 1983. Diel patterns of calling in nocturnal Orthoptera. Pages 45-72 in D. T. Gwynne and G. K. Morris (eds.): Orthopteran mating systems: sexual competition in a diverse group of insects. Westview Press, Boulder. Walker, T. J., and S. A. Wineriter. 1981. Marking techniques for recognizing individual insects. Fla. Entomol. 64:18-29. Walker, W. F. 1980. Sperm utilization strategies in nonsocial insects. Am. Nat. 115:780-799. Wiklund, C. 1982. Behavioural shift from courtship solicitation to mate avoidance in female ringlet butterflies ( Aphantopus hyperanthus ) after copulation. Anim. Behav. 30:790-793. Wing, S. R. 1982. Reproductive ecologies of three species of fireflies. M.S. Thesis, Univ. of Florida, Gainesville. NON-DIAPAUSE OVERWINTERING BY PIERIS RAPAE (LEPIDOPTERA: PIERIDAE) AND PAPILIO ZELICAON (LEPIDOPTERA: PAPILIONIDAE) IN CALIFORNIA: ADAPTIVENESS OF TYPE III DIAPAUSE-INDUCTION CURVES* By Arthur M. Shapiro Department of Zoology, University of California, Davis, California 95616 Diapause is generally regarded as a physiological adaptation which increases the probability of surviving the adverse season, and thus of reproducing after it is over. Many insect species show geographic differences in the environmental regimes which induce or inhibit diapause (e.g., critical photoperiod) and in the strength of the diapause induced. Such interpopulational differences are com- monly viewed as “fine tuning” to local climates, accomplished by natural selection and reflecting a genetic basis (e.g., Istock, 1981). Intrapopulational differences in photoperiodic sensitivity and dia- pause strength (e.g., chilling requirement) also occur, and have been interpreted as polymorphisms which “spread the risk” of environ- mental uncertainty over the population (cf. Bradshaw 1973, Shapiro 1979, 1980a). In multivoltine insects in seasonal climates, offspring produced by the last seasonal generation of adults are commonly induced to enter diapause by specific combinations of environ- mental factors; in mid-latitudes these are likely to be decreasing photophase/ increasing scotophase and decreasing or consistently low night temperatures. Warmer nights tend to shorten the critical photoperiod for a given population, or may effectively inhibit diapause altogether under field conditions. Beck (1980) characterized diapause induction as falling into four response types. Type I is the common long-day response, in which long days inhibit diapause and there is a single threshold beyond which diapause occurs. Type II, the short-day response, is the reciprocal of Type I. Type III has two well-defined critical day- lengths, such that diapause is induced between, say, 8 and 14 hours * Manuscript received by the editor April 15, 1984. 161 162 Psyche [Vol. 91 light, and inhibited when either less than 8 or more than 14 hr of light occur. Type IV is its reciprocal, in which diapause is inhibited over a relatively narrow range and induced both above and below. Of the four types, Type I is by far the commonest in temperate- latitude insects and Type IV the rarest. Low-latitude insects have rarely been examined for photoperiodic responses, though the different patterns of seasonality in low latitudes might be expected to produce a rather different picture than that seen in temperate zones. Type III curves are known from a variety of insects. Among Lepidopterans, two important species that show them are the European Corn Borer, Ostrinia nubilalis (Hbn.) (Beck, 1962) and the Large White, Pieris brassicae L. (Danilevskii, 1961). The adaptive significance of the short-day threshold is unclear. Nor- mally the entire population would have been induced by day-length- temperature interaction to enter diapause before the inhibition threshold was reached. Thus, the photosensitive stages would never be exposed to such short photoperiods in nature, and the existence of a short-day threshold would seem nonadaptive. This paper reports a situation in which Type III curves were demonstrably adaptive for two multivoltine Lepidopterans in nature. The Systems Papilio zelicaon Lucas and Pieris rapae L. are very common, widespread multivoltine butterflies in lowland California. Both have been monitored phenologically for up to 13 yr along a transect parallel to Interstate Highway 80 from sea level at the Suisun Marsh, Solano County, to treeline in the Sierra Nevada. We are concerned here with the phenology of populations at low elevations between the San Francisco Bay area and Sacramento. The flight seasons for both species may reach ten months (Table 1), with four (P. zelicaon) to about six (P. rapae ) generations per year. Both species have facultative pupal diapause under conventional photo- period-temperature control (Sims, 1980; Shapiro, unpublished data); both have Type III diapause-induction curves. Diapause is ir- reversibly determined no later than the fourth instar. P. zelicaon is the stronger diapauser and is unrecorded at any of our sampling sites between 1 December and 18 February. In some years the period with no rapae flying may, however, be as short as four weeks. Table 1. Dates of first and last observed flights of two common multivoltine butterflies in lowland north-central California, based on standardized methodology and level of effort. Pieris rapae Papilio zelicaon 1984] Shapiro — Pieris rapae 163 3 « W £ u. 03 03 O J c (U B 03 Ui o 03 CZJ w Ji Cfl « E Q X X X X X 22 JTj Os vn » ^ ^ OO cs! O — 0\^0'0'0'0'0'0' J3 1 1 03 _ 8 c c CS 3 o c u « o '5b — o O C 0 0 iS 73 o3 u T3 03 60 ^ C £ J compared to near-coastal populations. 164 Psyche [Vol. 91 In contrast, the Pierid Colias eury theme Bdv., which overwinters as a quiescent (non-diapausing) larva, flew continuously during the drought years of 1975-76 and 1976-77 and nearly so in 1983-84. Collections of mature larvae of both P. zelicaon and P. rapae made in late October and November over 12 yr have consistently given from 85-100% diapause pupae. Larvae of P. zelicaon are normally absent from all sites by the third week of November. Rapae larvae are occasionally found on garden cabbages and weedy Crucifers into early January. A fifth-instar larva was collected on a weed on 29 January 1979. The winter of 1982-83 produced 200% or more of normal rainfall over most of northern and central California. Nearly 20 cm of rain fell in March 1983 at our study sites, and the weather remained showery and unsettled into May. The spring flights of most butter- flies were late and poor; for many species the densities observed were the lowest in 12 yr. Both P. rapae and P. zelicaon were severely depressed at most sites; P. rapae was actually rare in spring, and the Willow Slough population of P. zelicaon, north of Davis, went extinct overwinter. The summer was unusually cool, cloudy, and moist. By autumn the populations of both species were near normal levels and at the Suisun Marsh P. zelicaon was commoner than usual at the end of the season. The latest flight recorded at Suisun for this species is 18 November (1973), a “false brood.” Although the species shut down nearly a month earlier in 1983, the autumn brood oviposited three to four weeks later than average. Similarly, at Davis, the last flight date of P. rapae was unexceptional but the population densities at the end of the flight were unusually high. For both species, these circumstances translated into an unprecedented number of larvae through mid-winter. In the Suisun Marsh, fourth- and fifth-instar larvae of P. zelicaon were common on Sweet Fennel, Foeniculum vulgare Mill. (Umbelli- ferae) on levees; collections were made on 28 December (4 L5, 1 L4, 1 L3) and 16 January (2 L5). In addition, one L3 was found on the same plant at Gates Canyon in the Vaca Hills, Inner Coast Range, near Vacaville, Solano County, 1 January. In the Davis area, a total of 67 larvae of P. rapae were collected from cultivated and weedy Cruciferae and Tropaeolum between 29 December and 15 January. Although L!-L3 were still numerous early in the period, only larger larvae (L4-L5) were taken. Numbers were so high that Cruciferous vegetables were seriously damaged in many gardens at this time. 1984] Shapiro — Pieris rapae 165 All larvae were placed in outdoor mesh cages over fresh cuttings of their hosts, and allowed to complete development under ambient conditions. The cages were less than 0.3 km from the U.C. Davis campus meteorological station. The first pupa of P. zelicaon was formed 13 January and the last 24 February. 50 of the 67 rapae larvae were parasitized by Apanteles (Cotesia) glomeratus (L.) (Hymenoptera: Braconidae) and failed to pupate; 5 died of un- known causes; 12 pupated between 9 January and 28 February. Results and Discussion Of the nine winter zelicaon, 1 prepupa died of unknown causes; 5 eclosed without diapause; and 3 apparently entered diapause. Eclosions occurred on 21 and 24 February and 1, 5, and 12 March 1984. As can be seen from Table 1, these dates coincide with the flight of P. zelicaon in the field. (At Gates Canyon, Vaca Hills, one zelicaon was found as early as 19 February 1984, the earliest flight ever recorded in the region.) All the butterflies which emerged were female, as was one of the 3 diapausers. Of the 12 healthy rapae pupae, all eclosed between 9 February and 24 March (seven males, five females). The flight began some- what earlier afield but continued throughout this period. The Apanteles wasps emerged without diapause in synchronized batches, each from its own host, between 24 January and 11 March — again, well-timed to parasitize first-instar larvae from eggs laid by first-brood rapae females. Diapause in this species is known to be under direct photoperiodic control (rather than mediated through the host, hormonally) in the U.S.S.R., but has not been studied in the introduced North American populations (Danilevskii, 1961). The phenotypes of emerging adults of both butterflies were compared with long series of field-collected specimens from Suisun and Davis from 1984 and prior years, including many individuals which must have come from diapaused pupae, and with reared ex- diapause individuals. No phenotypic differences which might permit the detection of non-diapaused adults in the spring populations were recognized. This result confirms experimental results which indicated that the post-diapause adult phenotype is not physio- logically coupled to the prior developmental arrest, but is rather a function of ambient conditions after reactivation (Shapiro, 1975a, 1978), superimposed on irreversible short-day prediapause deter- mination. 166 Psyche [Vol. 91 As is evident from Table 2, low-temperature lethality is a rare event in this part of California. Although diapause may confer a degree of frost-tolerance, its principal benefit in P. zelicaon and P. rapae in north-central lowland California seems to be to delay the onset of adult development until the bulk of the rainy season has passed; thus, adult eclosion coincides with sunny, warm weather suitable for flight and hence for reproduction. Pupae accumulate enough chilling by roughly mid-January to come out of diapause; subsequent development is timed by the weather, which thus determines not only the date of first flight but the degree of synchronization of the spring brood (Shapiro, unpublished data). The unusual 1983-84 winter larvae pupated at about the same time as normal diapausing pupae would have resumed development; hence it is not surprising that their subsequent development was synchronized with the wild population. (On the other hand, dia- paused pupae of Pieris napi L. ssp. will develop to the pharate adult at a constant temperature of 3°C, while non-diapause ones will not; hence in that species, diapausers might be expected to eclose first; Shapiro, unpublished.) The 3 diapausing zelicaon presumably cannot accumulate enough chilling to break diapause in spring 1984 and will thus lay over until early 1985. This would expose them to additional risks of mortality, and in terms of contribution to the rate of population increase, place them at a great selective disadvantage relative to nondiapausing members of their 1983-84 cohort (a delay of 4 generations). Multiple-year diapause occurs in most populations of P. zelicaon even under normal circumstances, although it is rare in the multi- voltine populations; it is especially common in univoltine foothill races which face unusually unpredictable and stressful climates and which would be expected to engage in “risk-spreading” (Sims, 1980). On the other hand, although pupae of P. rapae will remain viable for two years under constant refrigeration, no multiple-year dia- pause has ever been observed in that species under field conditions, and there is no indication that it has any physiological ability to diapause in summer (Shapiro 1975b, 1980b). Thus any diapause pupae produced by winter larvae of this species would presumably be doomed. In a Mediterranean climate, then, a Type III diapause-induction curve permits stragglers at the end of the season to complete development and enter the reproductive pool the following spring at Table 2. Climatic parameters for Sacramento (from U.S. Weather Bureau, NOAA Technical Memorandum NWS-WR-65, 1984] Shapiro — Pier is rapae 167 o £ O o ^ 3 oi Cu cd 3 rf -d- m © cn vo vO vO vo r" r- i^fNm(NooinvOd(N(N dO-mdO-<(NM- mvovovovot^i^t^r-r-' oooooooooo g" ■£ C n 2 3 "5 x) > o d Jr o o o o a> iu c/3 O O Z Z Q Q a -3 ^ £ 168 Psyche [Vol. 91 no disadvantage; it is a valuable hedge against off-season repro- duction. In such benign climates the photoperiodic threshold for diapause inhibition may be under intermittent selection at both ends of the curve. Non-diapause overwintering may be fairly frequent in some California butterflies, especially P. rapae. At the latitude of Sacramento it is definitely rare in P. zelicaon, since midwinter larvae were not observed in the first 12 yr of this study. In coastal southern California, however, this species flies more or less all winter (Emmel and Emmel, 1973; Orsak, 1977; Shapiro, unpub- lished) and larvae may often be found during the shortest days of the year. Although diapause intensity is reduced and thresholds are moderately shifted relative to further north, even the San Diego populations retain the ability to diapause (Sims, 1980). How often they use it under field conditions may depend on the timing of autumn reproduction on a year-by-year basis. The hypothesis that both thresholds of a Type III curve are adaptive and under selection is testable in principle by examining latitudinal shifts in critical photoperiods. We have had under study a culture of P. rapae from Xochimilco, D.F., Mexico, the southernmost (ca. 19° N) population of this species in the Americas; it also displays a Type III curve with the short-day inhibition threshold shifted upward (Shapiro, in prepara- tion). In climates where the onset of cold is more rapid than the change in photoperiod, that is, where strong air-mass contrasts exist and thermal lethality can occur with great suddenness, it is difficult to envision the low end of a Type III curve as adaptive. Cold nights should assure conservatism in the critical photoperiod, so that nearly all individuals will be determined as diapausers before the inhibition threshold can be reached. Larvae of P. rapae can be found into December and rarely into January at Philadelphia and New York City as well when early winter conditions are mild, but they never seem to survive. On the other hand, the occasional very early onset of lethal cold in such areas should select for more conservative diapause induction than the average conditions seem to warrant. The natural selection of diapause characteristics, already compli- cated by its nature as a recurrent (cyclical) process operating every 4th and 5th generation, is further complicated by the ability of stochastic variation to override it occasionally in benign climates. 1984] Shapiro — Pier is rapae 169 Acknowledgment This research is part of California Agricultural Experiment Station project CA-D*-AZO-3994-H, “Climatic Range Limitation of Phytophagous Lepidopterans,” AMS, principal investigator. References Beck, S. D. 1962. Photoperiodic induction of diapause in an insect. Biol. Bull. 122: 1-12. 1980. Insect Photoperiodism, 2nd ed. Academic Press,, New York. 387 pp. Bradshaw, W. E. 1973. Homeostasis and polymorphism in vernal development of Chaoborus americanus. Ecology 54: 1247-1259. Danilevskii, A. S. 1961. Photoperiodism and Seasonal Development of Insects. (English transl., 1965.) Oliver & Boyd, London, 238 pp. Emmel, T. C. and J. F. Emmel. 1973. The Butterflies of Southern California. Natural History Mus. of Los Angeles Co., Sci. Ser. 26. 148 pp. Istock, C. A. 1981. Natural selection and life history variation: theory plus lessons from a mosquito, in R. F. Denno and H. Dingle, eds., Insects Life History Patterns: Habitat and Geographic Variation. Springer-Verlag, New York. pp. 113-128. Orsak, L. J. 1977. The Butterflies of Orange County, California. Mus. Syst. Biol., U.C. Irvine, Res. Series 4. 349 pp. Shapiro, A. M. 1975a. Papilio “gothica” and the phenotypic plasticity of P. zelicaon (Papili- onidae). J. Lepid. Soc. 29: 79-84. 1975b. The role of watercress. Nasturtium officinale, as a host of native and introduced Pierid butterflies in California. J. Res. Lepid. 14: 158-168. 1978. The evolutionary significance of redundancy and variability in pheno- typic-induction mechanisms of Pierid butterflies (Lepidoptera). Psyche 85: 275-283. 1979. The phenology of Pieris napi microstriata (Lepidoptera: Pieridae) during and after the 1975-77 California drought, and its evolutionary significance. Psyche 86: 1-10. 1980a. Egg-load assessment and carryover diapause in Anthocharis (Pieridae). J. Lepid. Soc. 34: 307-315. 1980b. Mediterranean climate and butterfly migration: an overview of the California fauna. Atalanta 11: 181-188. Sims, S. R. 1980. Diapause dynamics and host-plant suitability of Papilio zelicaon (Lep- idoptera: Papilionidae). Amer. Midi. Nat. 103: 375-384. . CAMBRIDGE ENTOMOLOGICAL CLUB A regular meeting of the Club is held on the second Tuesday of each month October through May at 7:30 p.m. in Room 154, Biological Laboratories, Divinity Avenue, Cambridge. Entomolo- gists visiting the vicinity are cordially invited to attend. BACK VOLUMES OF PSYCHE Requests for information about back volumes of Psyche should be sent directly to the editor. F. M. Carpenter Editorial Office, Psyche 16 Divinity Avenue Cambridge, Mass. 02138 FOR SALE Reprints of artic les by W. M. Wheeler The Cambridge Entomological Club has for sale numerous reprints of Dr. Wheeler’s articles that were filed in his office at Harvard University at the time of his death in 1937. Included are about 12,700 individual reprints of 250 publications. The cost of the reprints has been set at 5c a page, including postage; for orders under $5 there will be an additional handling charge of 50> .£ u c .2 ■§ 2 3 S 2 3 -S £ "O Cl '5 fc 2 QJ (D 5 3 cd uj £ ■§ oo Jr .£ c ■a g j£‘ 00 *T c ■ — ti c/5 Jc ■- • — £■1 2 £ « £ 3 « •2 1 03 2 £ o C/3 CL 3 C ° _. «- •— rD 1/3 ~0 2 X) C 0- < D were 196 Psyche [Vol. 91 in amplexus upon our arrival in the field were still together several hours later when we departed. Pairs on flowers in the laboratory stayed together overnight. We found that these males were guarding females both prior to and following copulation (Dodson and Marshall 1984). Presumably time spent with one female, particularly after copulation has taken place, decreases a male’s opportunities for copulations with additional females. Why, then, might males be employing this guarding tactic? One possible answer is that the low population densities favor males that, upon finding a female, remain with her and actively repel other males. If such an attempt to monopolize a potential mate increased the chance of successfully copulating with her, precopulatory guarding might be a better tactic than spending this time searching for other females. This, however, seems an unlikely explanation for postcopulatory mate guarding. Parker (1970) argued persuasively that postcopulatory defense of a mate (preventing other males access to her) is an evolved strategy to counter sperm competition from potential, subsequent male mates. Although nothing is known regarding sperm precedence in ambush bugs specifically, most evidence for insects shows an insemination advantage for the last male to mate (Walker 1980). We feel there is another factor that may be important to our understanding of mate guarding in these ambush bugs (and perhaps in other insects). This factor is male preference for particular females. At one of our study sites, males were paired with females that were heavier than females without males (Dodson and Marshall 1984). These paired females also had more eggs than single females. The apparently superior fecundity of these females seemed to us to be a potential fitness gain for males. To further investigate the possible adaptiveness of selective pairing by males we looked at egg size within females. We had previously determined the wet weight (WW), pronotal width (PW) and femur length (FL) of these same females. All 3 measurements were significantly correlated with egg volume (BW: r = .60, p = .003; PW: r = .61, p = .003; FL: r = .62, p = .002). The 3 measurements are so highly correlated with each other, the residual variation examined by partial correlations was found to be negligible (rEv fl • ww pw = .04, p = .84; rEv pw • ww fl = .24, p = .3 1 ; rEV ww • pw fl — .27, p = .25). Therefore, all 3 variables are approximately equivalent predictors of egg volume. We believe. 1984] Dodson & Marshall — Ambush bug 197 as Johnson (1982) suggested, that this could potentially provide males with a basis for discriminating the most reproductively fit females. Males capable of assessing a cue to female fitness could “decide” whether to remain with a particular female or search for another. This would explain our findings of large females most often in pairs. Such a tactic could be even more feasible in insect populations which occur in high densities. The resulting high encounter rates would mean less time spent searching for more suitable mates. This hypothesis assumes that larger eggs enhance the success of resultant offspring (e.g., through increased nutrient provisions or greater competitive ability of larvae). Such advantages are contro- versial, but the rather limited data do yield some support. Relationships between egg size and offspring success in insects have been addressed by Capinera (1979 and references therein), Richards and Myers (1980) and Barbosa, et al. (1981). Discrimination of mates by males has been shown in several species in which males are contributing relatively large amounts of paternal care or other benefits (for insect examples see Thornhill and Alcock 1983). Perhaps more significant for this discussion are the findings of mate discrimination by males of noninvesting species (Loiselle; 1982; Verrel 1982; Hatziolos, M. E. and R. L. Caldwell 1983; references in Thornhill and Alcock 1983; Johnson and Hubbell, unpublished). Although some of the latter examples are laboratory experiments, the fact that the apparently adaptive behavior is exhibited suggests that selection under natural circum- stances must be occurring at significant levels. Several recent papers have reported overrepresentations of larger females within mating populations of insects. Such biases, based on either size or weight, have been found in beetles (McCauley and Wade 1978, Johnson 1982 and McLain 1982), pierid butterflies (Marshall 1982), ambush bugs (Dodson and Marshall 1984) and tephritid flies (Dodson, submitted). It would be interesting to determine the relationship between female body size and egg size in all such species in light of a male mate discrimination hypothesis. McCauley and Wade (1978) showed that mating male and female soldier beetles, Chaulignathus pennsvlvanicus, were heavier than nonmating males and females based on dry weights. They con- sidered dry weight to be an index of body size and suggested larger 198 Psyche [Vol. 91 females could be preferentially mating with larger (and presumably more fit) males by physically resisting mating attempts by all males and escaping the smaller ones. They further postulated that larger females might be “somewhat more receptive than small females”. However, except for the suggestion that receptivity might be related to egg development, they did not account for the paucity of small females in the mating population. Woodhead (1981) offered what appeared to be a more parsimonious explanation. She found that mating females had larger eggs than “rejecting” females and proposed that heavier female soldier beetles were more often found mating because they were more reproductively mature and therefore more receptive to mating. Woodhead (1981) did not discuss her choice of eggs for measurement. She did state that soldier beetles “mature eggs in groups” and so we assume that all primary oocytes were measured, including many not yet fully-yolked. Even if this were the case, however, her measurements are also consistent with the hypothesis that females differ in ultimate egg size and not just developmental stage. In many insects, oocytes are in various stages of development within the ovaries throughout most of a female’s adult life. This makes comparative measures of all eggs virtually impossible because of the miniscule size of the youngest oocytes. For valid comparisons between females, age classes of eggs must be de- lineated. Typically, fully-yolked eggs are chosen because they are easily defined, relatively easy to distinguish (although not always), and supposedly will not get any larger. We avoided this problem by measuring only chorionated eggs, i.e., eggs that are fully grown with their “shells” encasing them. Thus, we have demonstrated variation in egg sizes between females which are independent of sexual maturity and apparently dependent on female size. In summary, we have shown that P.fasciata males, when paired with females, aggressively fend off intruding males and that larger females produce larger eggs. Previous work (Dodson and Marshall 1984) revealed that males were more often paired with larger rather than small females in a natural population. All of these factors are consistent with the hypothesis that given some choice, males will preferentially guard mates which provide a higher reproductive potential. 1984] Dodson & Marshall — Ambush bug 199 Rl l ERENCES Barbosa, P., W. Cranksiiaw and J. A. Greenbi.att. 1981. Influence of food quantity and quality on polymorphic dispersal behaviors in the gypsy moth, Lymantria dispar. Can. J. Zool. 59: 293 296. Capinkra, J. L. 1979. Qualitative variation in plants and insects: effect of propagule size on ecological plasticity. Am. Nat. 114 (3): 350 361. Dodson, G. N. and L. D. Marshall. 1984. Mating patterns in an ambush bug, Phymata Jdsciala. (Phymatidae). Amer. Midi. Nat., in press. Hatzioi.os, M. E. and R. L. Caldwell. 1983. Role reversal in courtship in the stomatopod Pseudosqailla ciliata (Crustacea). Anim. Behav. 31: 1077 1087. Johnson, J. K. 1982. Sexual selection in a brentid weevil. Evolution 36 (2): 251-262. Loisli.i.k, P. V. 1932. Male spawning-partner preference in an arean-breeding teleost Cyprinodon mandarins californiensis Girard ( Atherinomorpha: Cypri- noclontidae). Am. Nat. 120 (6): 721 732. Marshall. L. D. 1982. Male courtship persistence in Colias philodice and C. eurytheme (Lepidoptera: Pieridae). J. Kans. Entomol. Soc. 55 (4): 729 736. McCauley, D. E. and M. J. Wade. 1978. Female choice and the mating structure of a natural population of the soldier beetle, Chauliognathus pennsylvanicus. Evolution 32: 771 775. McLain, D. K. 1982. Density dependent sexual selection and phenotypic assortative mating in natural populations of the soldier beetle, Chauliognalhus pennsylvanicus. Evolution 36 (6): 1227 1235. Richards, L. J. and J. H. Myers. 1980. Maternal influences on size and emergence time of the cinnabar moth. Can. J. Zool. 58: 1452 1457. Thornhill, R. and J. Ai.cock. 1983. The Evolution of Insect Mating Systems. Harvard University Press, Cambridge, MA. 547 pp. Verrel, P. A. 1982. Male newts prefer large females as mates. Anim. Behav. 30: 1254 1255. Woodiiead, A. P. 1981. Female dry weight and female choice in Chauliognathus pennsylvanicus. Evolution 35: 192 193. ON THE METAPLEURAL GLAND OF ANTS By Bert HOlldobler and Hiltrud Engel-Siegel Department of Organismic and Evolutionary Biology, MCZ-Laboratories, Harvard University Cambridge, Massachusetts Introduction The metapleural gland (also called metasternal or metathoracic gland), a complex glandular structure located at the posterolateral corners of he alitrunk is peculiar to the ants. Although the gland was noted by Meinert (1860) and Lubbock (1877), it was Janet (1898) who conducted the first detailed anatomical study of this organ, as part of his classic work on Mvrmica rubra. Additional details have been added by Tulloch (1936) on Mvrmica laevinodis; by Whelden (1957a, b, 1960, 1963) on Amblvopone ( Stigmatomma ) pallipes, Rhvtidoponera convexa, R. metallica, Eciton burchelli, E. ham- atum; by Tulloch et al (1962) on Myrmecia nigrocincta; and by Kiirschner* (1970) on Formica pratensis. It is generally assumed that the metapleural gland is a universal and phylogenetically old character of the Formicidae. Even the extinct species Sphecomyrma freyi of Cretaceous age appears to have possessed a metapleural gland (Wilson et al 1967a, b) and the organ is well developed in the most primitive living ant species Nothomyrmecia macrops (Taylor 1978) (see Fig. 2). In the course of our current comparative study of the internal and external anatomy of exocrine glands in ants, we discovered that the metapleural gland is absent or significantly reduced in several ant genera where such reduction had not been previously suspected. In addition we observed a widespread absence of the metapleural gland in males among ant species. Our survey is far from complete, even at the generic and tribal levels. We think, however, that the pattern revealed by our observations is important enough to warrant a short publication at this time. *Kurschner apparently was not aware that the paired thorax gland near the petiole she described was the metapleural gland. Manuscript received by the editor March 16, 1984. 201 202 Psyche [Vol. 91 Results The metapleural gland is a paired structure. Each side consists of a cluster of glandular cells, and each cell is drained via a duct into a membranous collecting sac that Tulloch et al. (1962) called secretory recess. The collecting sac leads directly into the storage chamber or atrium (receptacle, sensu Tulloch et al. 1962), a sclerotized cavity. Externally the metapleural gland is often marked by a pronounced vault (bulla), and a slit-shaped opening to the outside (Fig. 1). Although the metapleural gland is present in most ant species, it nevertheless varies greatly among them in size and shape (Figs. 2 & 3). Table 1 lists all of the ant species for which we obtained complete series of longitudinal sections through the mesosoma. The speci- mens were fixed in alcoholic Bouin or Carnoy, embedded in methylmethacrylate and sectioned 6 to 8 p thick. The staining was Azan (Heidenhain). We attempted to obtain approximations of the number of glandular cells either by counting the cells with clearly visible nuclei or by counting the number of duct openings in successive sections. The data reveal a considerable variation in the size of the metapleural gland among different species. Even more significantly, our study established that the gland is absent in Oecophvl/a /onginoda and O. smaragdina (Fig. 4), in all species of Camponotus and Po/vrhachis sectioned, and in Dendromyrmex chartifex (Tab. 1). We extended this list by an additional survey of the external features that indicate the presence of the metapleural gland, using light-stero-, and scanning electrone microscopy. Of 27 species of Camponotus investigated, only C. gigas showed a slit-shaped opening in the posterior metapleural region (Fig. 5c). In all other Camponotus species the metapleural gland is clearly absent (Fig. 5a, b; Tab. 2). This confirms the suggestion of Ayre and Blum (1971) based on external inspection of Camponotus pennsvlvanicus workers that this species might not possess a metapleural gland. In none of the species of Polyrhachis investigated did we detect any signs of a metapleural gland (Tab. 2). In addition our study revealed that in several species whose workers and queens have well- developed metapleural glands, the males do not possess this organ; whereas in other species the males have large metapleural glands (Tab. 1). 1984] Holldobler & Engel-Siegel — Metapleural gland 203 Fig. 1. The metapleural gland of Ana, illustrating the major anatomical feature of this organ, a. SEM micrograph of the mesosoma of A. cephalotes, showing the large pronounced vault (bulla) which covers the storage chamber. Arrow points to the slit-shaped opening (meatus), b. Longitudinal section through the mesosoma of A. se.xdens, showing the large , region of the metapleural gland (MPG). c. Longi- tudinal section through metapleural gland: CS = collecting sac; GC = glandular cells; M = meatus; R = storage chamber or receptacle. 204 Psyche [Vol. 91 Discussion The metapleural glands have been considered characteristic of all ants with the very few exceptions given by Brown (1968). From a survey of external criteria Brown listed four categories where the metapleural gland appears to be atrophied: “1. Males of army ants, subfamily Dorylinae. 2. Males of a few other genera, mainly in subfamily Myrmicinae (e.g. Leptothorax duloticus, Tetramorium, Strongylognathus, Rhoptromvrmex, H Liberia striata ). 3. Workers of the specialized slave makers of genus Polvergus. 4. Queens of certain scattered ant species that are known (or assumed, on grounds of other morphological peculiarities) to be social parasites, i.e., those species which found their colonies in the nests of other ant species”. From these findings Brown developed an intriguing hypothesis about the function of the metapleural gland: “the gland produces a substance that, when tasted or smelled, says to another ant colony, especially one of the same species, ‘I am an enemy’.” According to Brown’s hypothesis “an individual either with the same odor-or-taste, or with none at all, would be treated by its host colony as neutral”. This would explain why certain species whose individuals have to enter a foreign colony (social parasites; doryline males) often do not possess a metapleural gland. This hypothesis was challenged by Maschwitz (Maschwitz et al 1970, Maschwitz 1974). He was unable to experimentally demon- strate an enemy identification effect in the metapleural gland secretions, but he could show that in a number of ant species the metapleural gland secretions serve as powerful antiseptic substances that protect the body surface and nest against microorganisms. For example, the active antibiotic component of Atta sexdens was found to be phenylacetic acid, of which one ant stores an average of 1 .4 jug (Maschwitz et al 1970). In Crematogaster ( Phvsocrema ) difformis the secretions of the enlarged metapleural gland serve as antiseptics, but when discharged in larger quantities they can also repel animal enemies. Finally, in Crematogaster {Phvsocrema) inflata, which also possesses a hypertrophied metapleural gland, Maschwitz (1974) discovered that the sticky secretions function primarily as an alarm- defense substance. He hypothesized that in this case the antiseptic gland has evolved to become an alarm defense gland. Our discovery of the atrophy of the metapleural gland among more genera than previously suspected places this organ in a new 1984] Holldobler & Engel-Siegel — Metapleural gland 205 Fig. 2. Metapleural gland of a Nothomyrmecia macrops worker, a. SEM micrograph of meatus, b. Longitudinal section through the metapleural gland. CS = collecting sac; GC = glandular cells; R = storage chamber. 206 Psyche [Vol. 91 Fig. 3. (Above and facing page) Longitudinal sections through metapleural glands of workers of a. Myrmecia pilosula; b. Cerapachys ? turneri; c. Rhytidoponera melallica; d. Pseudomyrmex pallidus; e. Novomessor albiselosus; f. Myrmecocystus mendax. CS = collecting sac; GC = glandular cells; R = storage chamber. 1984] Holldobler & Engel-Siegel — Metapleura/ gland 207 208 Psyche [Vol. 91 light. Since the gland is absent in a number of ant species known to be extremely aggressive and discriminatory towards conspecific foreigners and interspecific competitors (for Oecophvlla see Holl- dobler and Wilson 1978, Holldobler 1979, 1984; for Camponotus see Carlin and Holldobler 1983) it is obvious that at least in these species the metapleural gland secretions have no function in enemy identification. The absence of the metapleural gland in male ants is also much more widespread than previously assumed. In fact, it appears that species in which males lack this organ or possess it in a very reduced state outnumber those in which the gland is well developed. Most of these males never have to enter a foreign colony in order to mate. Thus Brown’s argument concerning the absence of the metapleural gland in doryline males is further weakened. In our view Maschwitz’s experimental evidence concerning the antiseptic effect of most metapleural gland secretions is very convincing. We have repeatedly heard the argument that the secretions of other pheromone glands, such as the mandibular gland or poison gland are also acidic and have the potential of suppressing the growth of Escherichia coli in test plates. Thus, it is argued that Maschwitz’s tests, although demonstrating an antiseptic effect, do not necessarily prove a primarily antiseptic function of the meta- pleural gland. Maschwitz himself has pointed out that other exocrine glandular secretions frequently have antiseptic power. In fact, he hypothesized that most epidermal glands originally were antiseptic devices before they became more complex glandular structures that produce either repellent secretions against predators or alarm pheromones used in social communication (Maschwitz 1968, 1974; Maschwitz et al 1970). Metapleural gland secretions can freely flow out of the storage chamber. The meatus is sometimes densely covered with bristles (Fig. 6), and often there are hairs and dispenser bristles inside the atrium along which the secretion can easily flow to the outer surface (Fig. 6c). As Brown (1968) pointed out, “some ant species have been seen to draw the legs, especially the tibia and tarsi of the forelegs, repeatedly over the meatus of the gland and then rub these leg parts over the rest of the body”. In this way the metapleural gland secretion is probably spread over the whole body. It might also be distributed among nestmates by mutual grooming. Thus, it appears 1984] Holldobler & Engel-Siegel — Metapleural gland 209 Fig. 4. a. SEM micrograph of the mesosoma of a worker of Oecophylla longi- noda. b. Close-up of the posterolateral corners of the alitrunk. Arrows indicate the area where the opening of a metapleural gland should be located. 210 Psyche [Vol. 91 Fig. 5. (Above and facing page) SEM micrographs of the mesosoma of workers of several formicine species. Arrow indicates region where the opening of the metapleural gland should be located, a. Camponotus pennsylvanicus; b. Campo- notus consobrinus. In both species there are clusters of hairs visible in the area of the metapleural gland opening. Both species, however, lack a slit-shaped opening, c. The slit-shaped opening of the metapleural gland of Camponotus gigas. No opening can be detected in d. Colobopsis truncata; e. Dendromyrmex chart if ex; f. Polyrhachis ( Cyrtomyrma )? doddi. 1984] Holldobler & Engel-Siegel — Metapleura l gland 21 1 212 Psyche [Vol. 91 that the central location and general structure of the metapleural gland makes it ideally suited for distribution of an antiseptic secretion. Why then is the metapleural gland absent or strongly atrophied in Oecophylla, Polvrhachis, Dendromyrmex, most Camponotus, cer- tain social parasitic ants, and many male ants? Maschwitz et al (1970) offered the following explanations for the last two cases. Social parasitic ants, they argued, are usually highly attractive to the host ants, which groom them very frequently, so that the social parasites benefit from the social distribution of the antiseptic secretions of their host ants. This relieves the parasitic species of the burden of producing their own antiseptics and allows them to deploy the freed energy into other organs and functions. The absence of metapleural glands in male ants was given a different adaptive significance. Males live only a relatively short time inside the nest. They are also much less numerous than workers. Therefore, there exists no particular need for them to produce large amounts of antiseptic secretions. The latter hypothesis, of course, raises the question why in some species the males do have relatively large metapleural glands (Fig. 7f, Tab. 1 ). The reason could be that in those cases the ratio of males to workers might be much higher and/or the males might reside inside the nest for longer periods and therefore would present a considerable “antiseptic burden” to the colony. This would favor the selection of males capable of producing their own antiseptic secretions. Furthermore, the metapleural gland of male ants could also have another, secondary function, which does not exclude the primary antiseptic function; that is, it could produce sex phero- mones and hence be an important character maintained by sexual selection. During mating females might thus favor males with well developed metapleural glands, and the capacity to produce larger quantities of pheromone. It is interesting that in all weaver ant species studied the metapleural gland was atrophied. The species we checked included Oecophylla, Polvrhachis ( Cvrtomyrma )? doddi, Dendromyrmex, Camponotus xenex. It is reasonable to speculate that these arboreal ants are much less exposed to microorganisms than terrestrial ant species, and therefore an antiseptic metapleural gland became unnecessary. 1984] Holldobler & Engel-Siegel — Metapleural gland 213 Fig. 6. The metapleural gland of a worker of Iridomvrmex purpureus a. SEM micrograph of mesosoma. The arrow points to the opening of the metapleural gland, b. Longitudinal section through the metapleural gland, c. Close-up of a section through the glandular cells (GC) and dispenser bristles (B); D = glandular duct; R = collecting chamber. 214 Psyche [Vol. 91 Fig. 7. (Above and facing page) SEM micrographs of the exterior structures of the metapleural glands of several ant species, a. Alitrunk of Podomyrma pulchra worker, b. Close-up of bulla, slit-shaped opening and sensory hairs (?) of the metapleural gland of P. pulchra worker, c. Alitrunk of Crematogaster sp 10 (ANIC) worker, d. Alitrunk of Catalacus intrudens worker, e. metapleural gland opening with sensory hairs (?) of C. intrudens worker, f. Alitrunk of Crematogaster sp 10 (ANIC) male. Arrows point to the opening of the metapleural gland. 1984] HoHdobler & Engel-Siegel — Metapleural gland 215 Table 1. Evaluation of the histological preparations of the metapleural glands of ants. Species which could not be identified with certainty are listed either by the accession number of our histological collection, or by the species number assigned by the Australian National Insect Collection (AN1C), where voucher specimens have been deposited. metapleural gland present (+), absent (— ), and approximate 216 Psyche [Vol. 91 o W O o d as § I _ UJ < CT .C xi JS ^ 'E- o vi 2 -Si o E - s S ss ^ 1 1 < h D ® Q. 1 —5 CD CD [i] >2 < 5 Z U4 .2 2 § S5 I. 3 yj < £ Z o 5 & UJ z -s O 5 ft. T .g. <3 d § I S Leptogenvs nitida B. Holldobler, Shimba Hills, +(85) Kenya Leptogenvs regis B. Holldobler, Shimba Hills, + (98) Kenya 1984] Holldobler & Engel-Siegel — Metapleural gland 217 i i + + o o L M r oo £ w 22 3 I + o + ao uT 05 r , «3 2 c o E _o o CS X 3 C ed C/3 U 35 1 S 2 3 :0 < "D a £ 05 X jL> X O 2 o 1 ^ to .2 3 * < C/3 X 2 o 05 X T3 *J w| s > h- eg h- £ ■ W .to £ c/f £ 3 3 C/3 E X E z ea 05 3 05 X 3 C >. u u u. 4) 3 a. E X £ c T3 >> X c 3 u CZ! u C/3 u CQ ffl CO u- -q JU c u X o _o 2 .2 t/5 — x .2 o — 1 "5) 2 3 lu c 05 5 x ~ 2 5 4J X to X C 2 f— (/5 * 1 X c/5 ad —5 u ad ai -5’ "3 *1 .s CO * g ■§ | 5 ? | 1 .2 * 1 1 w "S’ "S' 0 ! > 5 .3 I' | 1 1 3 g 3 S 8 43 -s: -3 -3 -3 u ?3 S’ 6 5 5 5 -3' 1 ■3* > =s § 1 § W 3 < 3 ^ I -J -3 > 3 a: © O -g a 3 Neivamyrmex nigrescens B. Holldobler, Portal, +(201) Arizona, USA MYRMECIINAE B. Holldobler. Canberra, + (75) USA Philippine Rep. Queensland. Australia B. Holldobler. Eungella, + (38) Queensland. Australia B. Holldobler. Kuranda. +(137) Queensland, Australia B. Holldobler. Shimba Hills, + (85) B. Holldobler, Shimba Hills. + (98) Onychomyrmex hedleyi Onvchomyrmex sp. 2 (AN 1C) Onychomyrmex sp. I (AN 1C) Pachycondyla crassa DORYLINAE B. Holldobler. Port + (180) + (201) Holldobler & Engel-Siegel Table 1 . (Continued) Evaluation of the histological preparations of the metapleural glands of ants. Species which could not be identified with certainty are listed either by the accession number of our histological collection, or by the species number assigned by the Australian National Insect Collection (ANIC), where voucher specimens have been deposited. metapleural gland present (+), absent (— ), and approximate 218 Psyche [Vol. 91 "8 + t; £ + — o < •£ X E 03 "O c KJ 3 .■2 o ir co s_* CO U. U. C/3 cd s c >* , — jD o CD _o o C/3 15 CO 5 'n J _"S ’E o c CO o CO X o ffl U H DQ ui ad “3 ^ UJ * £ 5 Z r- U -s; S3 S3 V O O Novomessor albisetosus B. Holldobler, Portal, Arizona, +(250) USA Novomessor cocker e Hi B. Holldobler, Portal, Arizona, +(272) +(62) USA 1984] Holldobler & Engel-Siegel — Metapleura/ gland 219 i o On + oc r~- + ON rn -'-t I ON • • — ^ + 2 4- G. + I < C/2 ad ui ad ad d < C/3 CO C a 3 xz C8 X r 3 c OJ C/3 >> o c N c X c o CO CO CJ CO CL C/2 u < u o E uT ^ CO > JD .2 JJ . 03 o 2 0 1 75 •_ C/3 3 < o T3 X X 1 2 o 1 X <£ o =5 X .2 CO > CO oc cd < cd cd CQ cd "p I -c UJ < 3 Z p u s uj ~ 2 * O R- x S H -S O o Z ^ UJ -R < $ Z ~ < T O' 3 UJ CJ -C 3 UJ p < ~ z S u | 2 2 Q£. r" O r* U. X X X o o o ■a T3 zz < ~ < za ■■o C/2 :0 C/2 :0 X X X X X CQ ad ad _ea ao 3 O Q o CL JL> X - o 5 s 5' a C3 s § § Q k. U, -s: CL [Vol. 91 I Qld, Australia Polvergus rufescens B. Holldobler, Gerbrunn, Bavaria, Germany Polyrhachis (Cyrtomyrma) B. Holldobler, Port Douglas, ? doddi Qld, Australia 1984] Holldobler & Engel-Siegel — Metapleura/ gland 221 Table 2. An external survey of the presence (+) or absence (— ) of features that indicate a metapleural gland in the genera Camponotus, Dendromyrmex, and Polyrhachis. Further information concerning species identification see Table 1. species metapleural collector and locality gland Camponotus abdominalis floridanus N. Carlin, Sugarloaf Key, Florida USA C. ? aeneopilosus (sp. 3, ANIC) B. Holldobler, Canberra, Australia C. americanus N. Carlin, Blue Hills, Massachusetts, USA C. castaneus P. Calabi, Tall Timbers, Florida, USA C. ? ephippium (sp. 13, ANIC) B. Holldobler, Canberra, Australia C. ferrugineus N. Carlin, Blue Hills, Massachusetts, USA C. gigas ?, S-Sumatra + C. herculeanus B. Holldobler, Gramschatz, Bavaria, Germany C. ? intrepidus (sp. 4, ANIC) B. Holldobler, Canberra, Australia C. midas (acc. #190) B. Holldobler, Poochera, Australia C. nearcticus N. Carlin, Blue Hills, Massachusetts, USA C. ? nigriceps (sp. 2, ANIC) B. Holldobler, Poochera, Australia C. perthiana B. Holldobler, Canberra, Australia C. planatus B. Holldobler, Keys, Florida, USA C. schaeferi M. Moglich, Portal, Arizona, USA C. senex W. M. Wheeler, Pueblo Nuervo, Panama C. sericeiventris J. Traniello, BCI, Panama C. socius B. Holldobler, Tampa, Florida, USA C. ? suffusus (acc. #236) B. Holldobler, Canberra, Australia C. tortuganus N. Carlin, Sugarloaf Key, Florida 222 Psyche [Vol. 91 Table 2. (Continued) An external survey of the presence (+) or absence (— ) of features that indicate a metapleural gland in the genera Camponotus, Dendro- nivrmex, and Polyrhachis. Further information concerning species identification see Table 1. species collector and locality metapleural gland C. vicinus N. Carlin, N. Franks, Cortez, Colorado, USA C. sp. 10 (ANIC) B. Holldobler, Poochera, S. Australia , 1 B C. sp. 14 (ANIC) B. Holldobler, Canberra, Australia C. sp. 19 (ANIC) (walkeri group) B. Holldobler, Cooktown, Qld, Australia - C. sp. 20 (ANIC) B. Holldobler, Canberra, Australia - C. sp. 21 (ANIC) B. Holldobler, Kuranda, Australia - C. sp. 22 (ANIC) (clareipes group) B. Holldobler, Bateman’s Bay, NSW, Australia - Dendromvrmex fabricii J. Wenzel, BCI, Panama - Polyrhachis ? amnion (acc. #65) B. Holldobler, Bateman’s Bay, NSW, Australia - P. 1 gab (acc. #102) B. Holldobler, Mt. Carbine, Qld, Australia - P. schlueteri B. Holldobler, Shimba Hills, Kenya - P. ? sokolova (acc. #95) B. Holldobler, Port Douglas, Qld, Australia - P. sp. 2 (ANIC) B. Holldobler, Canberra, Australia - P. sp. 3 (ANIC) B. Holldobler, Canberra, Australia - On the other hand other arboreal species, such as Pseudo- myrmex, Podomyrma, Catalaucus (Fig. 7), Opisthopsis respiciens, and Lasius fuliginosus have well-developed metapleural glands, whereas some terrestrial Polyrhachis and Camponotus have lost their metapleural gland. But this does not contradict our specula- tion. An arboreal life style might favor but not necessitate a reduction and atrophy of the metapleural gland.* Furthermore, most Camponotus and Polyrhachis species are arboreal or live in or *In a similar manner, the social parasitic life style appears to favor but does not necessitate the reduction of the metapleural gland (see Maschwitz. et al 1970). 1984] Hoddobler & Engel-Siegel — Metapleural gland 223 on large pieces of dead wood. It is entirely possible that con- temporary terrestrial species might have originiated from arboreal ancestors that had already lost their metapleural gland. We have currently no explanation, however, why Camponotus gigas appears to have retained or redeveloped its metapleural gland. Of course, it would also be interesting to known how those terrestrial species that have lost their metapleural gland defend themselves and their nests against microbial and fungal attacks. Acknowledgements We thank all the collectors mentioned in Table 1 and 2, and B. Bolton, W. L. Brown, R. Snelling, R. W. Taylor for helping us with the identification of many species, and Ed Seling for his assistance during the SEM work. Special thanks to R. W. Taylor and the Division of Entomology, CSIRO, Canberra (Australia) who in many ways supported this study. The work was further supported by grants from the National Science Foundation, the National Geographic Society and the John Simon Guggenheim Foundation. Appendix We would like to take this opportunity to correct some errors which appeared in two of our previous publications on exocrine glands in ants. 1) In our paper “Tergal and sternal glands in ants” (Psyche 85, 285-330, 1978) on page 297, (Table la), Myrmecinae should be corrected to Myrmicinae; on page 298, the locality of Veromessor pergandei, given as Mexico, should be corrected to Arizona; on page 299, the species PaChycondyla spec, listed under the subfamily Formicinae should be corrected to Acantholepis spec. 2) In our paper “Tergal and sternal glands in male ants” (Psyche 89 , 1 13-132, 1982), Fig 1 B, the lettering PG should be changed to P, because it indicates part of the penis with the penis gland, and not the pygidial gland (PG), shown in Fig. 1A. References Ayre, G. L. and M. S. Bu m. 1971 Attraction and alarm of ants ( Camponotus spp.— Hymenoptera: Formicidae) by pheromones. Physiological Zoology 44: 77-83. 224 Psyche [Vol. 91 Brown, W. L. 1968. An hypothesis concerning the function of the metapleural glands in ants. Amer. Nat. 102: 188- 191. Carlin, N. F. and B. HOi.ldobi.hr. 1983. Nestmate and kin recognition in interspecific mixed colonies of ants. Science 222: 1027 -1029. H0li.dobi.hr, B. 1979. Territories of the African weaver ant ( Oecophylla longi- noc/a [Latreille] ). A field study. Z. Tierpsychol. 51: 201-213. HOlldobi.hr, B. 1983. Territorial behavior in the green tree ant ( Oecophylla smaragdina). Biotropica, 15: 241-250. HOi.ldobi.hr, B. and E. O. Wilson. 1978. The multiple recruitment system in the African weaver ant Oecophylla longinoda (Latreille) (Hymenoptera: Formi- cidae). Behav. Ecol. Sociobiol. 3: 19-60. Janht, Cil 1898. Etudes sur les Fourmis, les Guepes et les Abeilles, Note 17: Systeme glandulaire tegumentaire de la Myrmica rubra. Observations diverses sur les Fourmis. Paris, George Carre et C. Naud, Editeurs, pp. 1-30. KCrschnhr, I. 1970. Zur Anatomie von Formica pratensis, Retzius, 1783. Beitr. Ent. 20: 375-387. Lubbock, J. 1877. On some points on the anatomy of ants. The Monthly microsc. Journal 18: 121 142. Maschwitz, U. 1969. Wehrdriisensysteme und Wehrverhalten bei Dytisciden. Verh. Dtsch. Zool. Ges. Insbruck (1968) Zool. Anz. 32(suppl): 410-416. Masc hwitz, U. 1974. Vergleichende Untersuchungen zur Funktion der Amei- senmetathorakaldriise. Oecologia 16: 303-310. Maschwitz, U., K. Koob und H. Schildknhcht. 1970. Ein Beitrag zur Funktion der Metapleuraldriise der Ameisen. J. Ins. Physiol. 16: 387-404. Mhtnhrt, F. 1860. Bidrag til de danske Myrers Naturhistorie. Dansk Vetensk. Selskab. 5: 275-340. Taylor, R. W. 1978. Nothomyrmecia macrops: a living-fossil ant rediscovered. Science 201: 979 985. Tulloc h, G. S. 1936. The metasternal glands of the ant Myrmica rubra, with special reference to the golgi bodies and the intracellular canaliculi. Ann. Entomol. Soc. Amer. 29: 81-84. Ti t i ck h, G. S., J. E. Shapiro, and B. Hhrshhnov. 1962. The ultrastructure of the metasternal glands of ants. Brooklyn Entomol. Soc. Bull. 77: 91-101. Whhldhn, R. M. 1957a. Notes on the anatomy of the Formicidae 1. Stigma- tomma pallipes ( Haldeman). J. New York Ent. Soc. 65: 1-21. Whhldhn, R. M. 1957b. Notes on the anatomy of Rhytidoponera convexa Mayr (“violacea” Forel) (Hymenoptera: Formicidae). Ann. Ent. Soc. Amer. 50: 271-282. Whhldhn, R. M. 1960. The anatomy of Rhytidoponera metallica F. Smith (Hymenoptera: Formicidae). Ann. Ent. Soc. Amer. 53: 793-808. Whhldhn. R. M. 1963. The anatomy of the adult queen and workers of the army ants Eciton burchelli Westwood and Eciton hamatum Fabricius. J. New York Ent. Soc. 71: 90 1 15. Wilson, E. O., F. M. Carphntkr and W. L. Brown. 1967a. The first Mesozoic ants, with the description of a new subfamily. Psyche (Cambridge) 74: 1-19. Wilson, E. O., F. M. Carpknthr and W. L. Brow n. 1967b. The first Mesozoic ants. Science 157: 1038-1040. A NEW EXOCRINE GLAND IN THE SLAVE RAIDING ANT GENUS POLY ERG US By Bert HOlldobler* Department of Organismic and Evolutionary Biology, MCZ-Laboratories, Harvard University, Cambridge, Massachusetts Introduction In his classical study of the workers and males of Myrmica rubra, Janet (1898, 1901) discovered a pair of clusters of a few glandular cells under the 6th abdominal tergite. Each cell is drained by a duct that penetrates the intersegmental membrane between the 6th and 7th abdominal tergites. Recent investigations have demonstrated that this gland is very common in ants, but it varies considerably in size and structure (Holldobler and Engel 1978). Since the gland is anatomically closely associated with the last exposed tergite in female ants (7th abdominal tergite = pygidium) Kugler (1978) sug- gested that it be called the pygidial gland. Recently Jessen and Maschwitz (1983) proposed the alternative name, Janet’s gland, in honor of its discoverer Charles Janet. The pygidial gland has been found in workers of representative species belonging to all subfamilies except the Formicinae (for liter- ature review see Holldobler and Engel 1978; Holldobler and Engel- Siegel 1982; Holldobler 1982). I report here the first discovery of a pygidial gland in Poiyergus, a genus of the subfamily Formicinae. But the anatomy has unusual features that point to a possibly indepedent origin in evolution. Material and Methods Three species of Poiyergus were investigated. The European spe- cies Poiyergus rufescens was collected near Wurzburg (W-Ger- many), the two North American species P. breviceps and P. lueidus were found near Portal (Arizona) and near Rocky Point in Suffolk County (New York), respectively. Virgin queens and males were * Manuscript received by the editor March 24, 1984. 225 226 Psyche [Vol. 91 available only in the case of P. lucidus. For comparison two com- mon slave species of P. rufescens, Formica rufibarbis and F. fusca (from the Wurzburg area), as well as two species of the slave raiding Formica sanguinea group (collected in Massachusetts), were also studied. For histological investigations life specimens were fixed in alco- holic Bouin or Carnoy (Romeis 1948), embedded in methylmetha- crylate, and sectioned 6-8 p thick with a D-profile steel knife on a Jung Tetrander I microtome (Rathmayer 1962). The staining was Azan (Heidenhain). The SEM pictures were taken with an AMR 1000 A scanning electron microscope. Results Workers and queens of the three Polyergus species investigated possess a large, tergal complex gland. One part of it is located between the 6th and 7th abdominal tergites; the other part belongs to the 7th tergite (Fig. 1). Because of its anatomical location I pro- pose tentatively to call this organ pygidial gland, although it differs in several respects from the pygidial glands found in other ant subfamilies. t The pygidial gland of Polyergus consists of a reservoir, formed by an invagination of the intersegmental membrane between 6th and 7th abdominal tergites. Glandular cells (~80 cells in P. rufescens workers) are drained into this reservoir through ducts penetrating the intersegmental membrane. Ducts of a second group of glandular cells open into a series of cuticular cups located along the anterior margin of the 7th abdominal tergite (Fig. 1 , 2). This striking cuticu- lar structure is usually not visible, because it is covered by the poste- rior portion of the 6th abdominal tergite. It is, however, easily exposed by slightly pulling the two terminal abdominal tergites apart. Although only a few specimens of each species were investi- gated in detail, species-specific differences in the pygidial cup struc- ture are so conspicuous that this organ might be considered a valid taxonomic character. The cup structure is most strikingly developed in workers of P. rufescens (Fig. 2). In this case several cups are often merged to form one large cup containing 2-5 glandular cell openings. I counted a total of approximately 250 duct openings on the pygidium of P. rufescens workers. In workers of P. breviceps the cup structure is 1984] Hdlldobler — New exocrine gland 227 Fig. 1. Above: Schematical illustration of the Polvergus pygidial gland between 6th and 7th abdominal tergites. Below: Longitudinal section through the pygidial gland of a P. rufescens worker. CS: cup structure; CU: cup; GC: glandular cells; R: intersegmental reservoir. 228 Psyche [Vol. 91 Fig. 2. SEM-photographs of the pygidial cup structure of a P. rufescens worker. Above: overview of the pygidium (= 7th abdominal tergite). Below: close-up of the cup structure. Note the duct openings inside the cups. 1984] Holldobler — New exocrine gland 229 smaller (with a total of ~ 150 duct openings), but in this species as well several small cups frequently form one large cup (Fig. 3). In P. lucidus workers the pygidial cup structure is still smaller, with approximately 100 duct openings. In this case the cup structure consists primarily of single cups (Fig. 4). It is interesting to note, however, that in P. lucidus queens the pygidial cup structure is considerably larger (~200 duct openings), and many of the cups are merged (Fig. 5). External morphological studies of P. lucidus males indicate that they lack the organ. Formica rufibarbis and F. fusca, two common slave species of P. rufcscens, as well as two representatives of the slave raiding Formica sanguinea group were found not to possess a pygidial gland (Fig. 6). Discussion Of a total of 17 formicine species investigated (the current study and Holldobler and Engel 1978) belonging to 6 genera ( Acanthole - pis, Camponotus, Formica, Myrmecocystus, Oecophylla, Polyer- gus) only those in Polvcrgus possess the pygidial gland. This structure is a complex gland with one group of glandular cells open- ing into an intersegmental reservoir, and another group of cells being drained into cuticular cups in the 7th abdominal tergite. From these features and from its general anatomy I conclude that the pygidial gland of Polyergus is not homologous with the pygidial gland of other ant subfamilies, but has evolved independently, pos- sibly in connection with the highly specialized slave raiding behavior of Polyergus . Polyergus slave raids are organized by scout ants that deposit chemical trails (Talbot 1967). The precise origin of the trail phero- mones has not yet been established. In most formicine species secre- tions from the rectal bladder serve as trail pheromones (for review see Holldobler 1978). These function in many cases primarily as orientation cues and are often supplemented by excitement (or arousal) signals, which can be chemical or mechanical in nature. Recent observations suggest that recruitment to slave raids in Polyergus might be based on a similar mechanism. Topoff et al. (1984) found that individual scout ants lead the raiding party of nestmates to the slave ants’ nest. He observed that the ants, although following a chemical trail, tend to “swarm” 230 Psyche [Vol. 91 Fig. 3. SEM-photograph of the pygidial cup structure of a P. breviceps worker. Above: overview of the pygidium. Below: close-up of the cup structure. 1984] Holldobler — New exocrine gland 231 Fig. 4. SEM-photograph of the pygidial cup structure of a P. lucidus worker. Above: overview of the pygidium. Below: close-up of cup structure. 232 Psyche [Vol. 91 Fig. 5. SEM-photograph of the pygidial cup structure of a P. lucidus queen. Above: overview of the cup structure at the anterior margin of the 7th abdominal tergite. Below: close-up of the cup structure. Arrows point ot the intersegmental duct openings. 1984] Holldobler — New exocrine gland 233 Fig. 6. SEM-photograph of the pygidium of a worker belonging to the Formica sanguinea group. Note: no cup structure on the anterior margin of the 7th abdominal tergite. around the leader ant, often moving briefly ahead, but always returning to the leader. Myrmecologists have long noted this pecu- liar looping behavior at the front of the raiding columns of Poly- ergus (see for example Schmitz 1906), but only the recent studies by Topoff have indicated that this appears to be due to the orgainizing role and excitement emanating from the leader ant. These observa- tions suggest that the leader discharges not only a longer lasting trail pheromone but also an arousal signal, possibly from the pygidial gland. Similar recruitment patterns have been observed in Camponotus socius and Myrmecocystus mimicus, where secretions from the poi- son gland seem to function as arousal signals (Holldobler 1971, 1981), as well as in Camponotus ephippium, where the arousal sig- nal appears to originate from the cloacal gland (Holldobler 1982). However, this hypothesis cannot be applied to the well developed pygidial gland in Polyergus queens. Wasmann (1915) first proposed that a young Polyergus queen, in order to found a new colony, has to intrude a colony of a slave ant species, where she kills the resident 234 Psyche [Vol. 91 queen and is subsequently adopted as the “replacement” queen by the slave ant workers. This mode of parasitic colony founding was later confirmed in field observations by Gosswald (1932). It is possible that the Polvergus queen achieves her acceptance in the foreign colony by emitting either a supernormal attractant or a disorganizing alarm substance (hence, some form of queen “propa- ganda” substance), possibly discharged from the pygidial gland. In fact, the pygidial gland secretions of Polyergus workers might also function as a kind of propaganda allomone when encountered by slave ants, very similar to the Dufour’s gland secretions of the slave raiding species Formica pergandei and Formica subintegra which have a confusing propaganda effect on the raided slave ant colony (Regnier and Wilson 1971). Further experimental work is required to determine which, if apy, of these proposed speculations on the function of the Polyergus pygidial gland is correct. Acknowledgements I thank Hiltrud Engel-Siegel and Ed Seling for technical assist- ance, and Howard Topoff and Linda Goodloe who generously pro- vided specimens of Polyergus breviceps and P. lucidus. This work was supported by NSF grant NBS 82 19060. References GOsswald, K. 1932. Okologische Studien iiber die Ameisenfauna des mittleren Maingebietes. Zeitschrift f. Wissensch. Zoologie 142: 1-156. HOlldobler, B. 1971. Recruitment behavior in Camponotus socius (Formicidae). Z. vergl. Physiol. 75: 123-142. HOlldobler, B. 1978. Ethological aspects of chemical communication in ants. Advances in the Study of Behavior. 8: 75-1 15. HOlldobler. B. 1981. Foraging and spatiotemporal territories in the honey ant Mvrmecocys- tus mimicus Wheeler (Hymenoptera: Formicidae). Behav. Ecol. Socio- biol. 9: 301-314. HOlldobler, B. 1982a. Chemical communication in ants: new exocrine glands and their behav- ioral function. In: The Biology of Social Insects (ed. M. Breed, C. D. Michener, H. E. Evans). Westview Press, Boulder, Colorado. 1984] Holldobler — New exocrine gland 235 HOlldobler, B. 1982b. The cloacal gland, a new pheromone gland in ants. Naturwissenschaften 69: 186. HOlldobler, B. and H. Engel 1978. Tergal and sternal glands in ants. Psyche (Cambridge) 85: 285-330. HOlldobler, B. and H. Engel-Siegel 1982. Tergal and sternal glands in male ants. Psyche (Cambridge) 89: 1 13-132. Janet, Ch. 1898. Etudes sur les Fourmis, les Guepes et les Abeilles, Note 17: Systeme glandulaire tegumentaire de la Myrmica rubra. Observations diverses sur les Fourmis. Paris. Georges Carre et C. Naud, Editeurs pp. 1-30. Janet, Ch. 1902. Anatomie du gaster de la Myrmica rubra. Paris, Georges Carre et C. Naud, Editeurs pp. 1-63. Jessen, K. and U. Maschwitz 1983. Abdominaldrusen bei Pachvcondyla tridentaia (Smith) (Formicidae, Ponerinae). Insectes Sociaux 30: 123-133. Khgler, Ch. 1978. Pygidial glands in myrmicine ants (Hymenoptera, Formicidae). Insectes Sociaux 25: 267 274. Rathmayer, W. 1962. Methylmetacrylat als Einbettungsmedium fur Insekten. Experientia (Basel) 18: 47-48. Regnier, F. E. and E. O. Wilson 1971. Chemical communication and “propaganda” in slave maker ants. Science 172: 267-269. Romeis, B. 1948. Mikroskopische Technik. Miinchen 1948. Schmitz, H. 1906. Das Leben der Ameisen und ihrer Gaste. Verlagsanstalt vorm. G. J. Manz, Buch-u. Kunstdruckerei, Miinchen-Regensburg. Talbot, M. 1967. Slave raids of the ant Polyergus lucidus Mayr. Psyche (Cambridge) 74: 299-313. Topoff, H., B. LaMon, L. Goodloe, and M. Goldstein 1984. Social and orientation behavior of Polyergus breviceps during slave- making raids. Behav. Ecol. Sociobiol. 15: 273-279. Wasmann, E. 1915. Gessellschaftsleben der Ameisen, Munster, 1915. COLONY COMPOSITION IN FOUR SPECIES OF POLISTINAE FROM SURINAME, WITH A DESCRIPTION OF THE LARVA OF BRACHYGASTRA SCUTELLARIS (HYMENOPTERA: VESPIDAE) By James M. Carpenter* and Kenneth G. Ross** Introduction Knowledge of the social structure in colonies of tropical Polisti- nae remains fragmentary. Since the pioneering studies of Richards and Richards (1951), capture and dissection of the adult population of a nest has been the primary method for reconstruction of the caste composition of a species. As the reviews of Jeanne (1980) and Akre (1982) document, long-term observational studies are increas- ingly contributing to behavioral analysis in this group. Yet the pre- vailing view of social structure in swarm-founding Polistinae is still largely based upon limited studies of a few species. Indeed, most of the species studied by Richards and Richards (1951) have not been re-examined. While participating in the Second Cornell Suriname Expedition, the senior author collected five colonies of three of the species stud- ied by Richards and Richards (1951), and one species studied by Richards (1978). Dissection of the specimens revealed certain dis- crepancies with the published accounts of colony composition for these species, as well as new information. These results are presented below, together with descriptive notes on the nests. The larvae of Polybia bistriata, P. catillifex and Metapolybia cingulata have been described by Reid (1942). He also described the larva of a Brachy - gastra species, termed by him Nectarinia scut el laris. However, Richards (1978: 166) stated that the nests he had obtained in Guyana, upon which Reid based his work, were actually those of the species myersi. The larva of B. scutellaris has thus never been de- scribed. A description is included in this paper. *Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138. **Department of Entomology, University of Georgia, Athens, Georgia 30602 Manuscript received by the editor May 11, 1984. 237 238 Psyche [Vol. 91 Methods The colonies were collected during February 1982 in the Raleigh Vallen-Voltzberg Natuureservaat (Foengoe Eiland 4°42'N, 56° 12"W, approximately 90 m elev.; Voltzberg Camp 4°42'N, 56° 13'W, also 90m), Saramacca Dist., Suriname. The nests were placed in plastic bags and the adults killed with ’’wasp freeze”. Adults and larvae were both preserved in Dietrich’s fluid until dissection. All female wasps were dissected and various measurement data taken with an optical micrometer. Length of the discal cell (Ml) was used as a measure of wing length; the left wing was excised and flattened in a drop of glycerin for measurement. No notable wing wear was found for any of the wasps. A modified Cumber ovariole index (Cumber, 1949) was used to quantify ovarian development, with an additional count of oocytes greater than 0.6 mm in length (considered “deve- loped”) as a check. Inseminated individuals could readily be distin- guished by the pronounced reddish-golden color of the spermatheca; nevertheless all spermathecae were examined under phase-contrast microscope. Sperm were visible as an agglutinated mass in the lumen (Fig. 1). Classification into castes separated queens (insemi- nated individuals), intermediates (uninseminated but with an ovari- ole index of 1 or more) and workers (uninseminated, index = 0). Richards’ gland (on metasomal sternum V) was present in all species (Fig. 2). No caste differences were observed in this structure. Exploratory data analysis (Velleman and Hoaglin, 1981) was per- formed, with further analysis where results indicated. Analyses included investigations of the relationship between the various mor- phometric measures (were the correlations positive, and so reflec- tions of general size?), the relationship between these and ovarian development (is size associated with reproductive capacity?), and differences between castes in these measurements. The data were normally distributed, therefore the results of t-tests are reported for caste differences. Larvae were examined with the aid of cuticular stains (acid fuch- sin, Grenacher’s alcoholic borax-carmine, eosin-Y). Mouthparts and the frontoclypeal region of B. scutellaris were excised, and the latter examined under phase-contrast miscroscope. Voucher specimens of all species have been deposited in the Cor- nell University Insect Collection, Lot number 1 129. 1984] Carpenter & Ross — Polistinae 239 Fig. I. Brachygasna scute/laris, spermatheca with sperm in lumen. Results Polvbia bistriata: Two colonies of this species were collected on Foengoe Eiland on February 4. The nests were located within 4 m of one another on the undersides of leaves in shaded secondary growth along a path. Both were about 1 .5 m above the ground and thus inconspicuous. Several adults escaped from each nest. The nests were similar in appearance, and of the same shape and size as depicted in Figs. 19-21 of Richards and Richards (1951). The envelope was whitish-gray with brown streaks, and moderately flex- ible. Under high magnification, fragments of plant tissue were dis- tinguishable among masticated, somewhat granular material. The envelopes were built by the “edge technique” (Jeanne, 1973), and were not thickened. They were evidently the products of single con- struction efforts, and no growth lines were present. The entrances were slightly off-center and ventral, unlike Fig. 19 of Richards and Richards but like those described in the text (p. 56). 240 Psyche [Vol. 91 Fig. 2. Polybia catillifex, Richard’s gland (700X), ventral view. The first colony (82-02-04-1) contained a single comb of 147 uncapped cells, 24 of which contained early-instar larvae and 48 with eggs. These were concentrated in the center of the comb. No sign of cell re-use was present; the nest appeared to be quite young. Twenty-six adults were captured, six of which proved to be queens. Among the 20 workers, one had developing oocytes in the ovarioles, but these were still smaller than the accompanying trophocyte masses. Measurements consisted of maximum apical width of meta- somal tergum I and length of this segment taken from the ligament base to the apex, as well as length of the discal cell. The tergal measurements were not significantly correlated with each other (all N=26; r=.303, Pearson product-moment correlation), but were correlated with length of the discal cell (r=.482, p<.05 for tergal width; r =.488, p <.01 for length). Hamulus number was not signifi- cantly correlated with any measurement (r=.063 for tergal width, .076 for length, and .157 for discal cell length). Only discal cell length was significantly correlated, negatively, with ovariole index 1984] Carpenter & Ross — Polistinae 241 (r = — .443, p<.05), however, the trends were also negative for the other measurements (r = — .307 for tergal width, —.338 for length, and —.050 for hamulus number). Queens did not differ significantly from workers in tergal width, discal cell length or hamulus number (t = — 1 .883, —1.917, and 0.070, respectively). Queens had signifi- cantly shorter terga than workers when compared by a t-test (mean = 1 .47 mm versus 1 .52, t = -2.470, p < .03). The second colony (82-02-04-2) had a comb of 151 cells. Of these, 25 were capped, and some contained partly melanized pupae. There were also late-instar larvae in some uncapped cells; the number could not be ascertained due to damage to the nest, but was not greater than 12. Other cells were empty. No sign of cell re-use was present, and the occupied cells were irregularly arranged but mostly central. This colony was probably also producing its first brood. Forty-six adult wasps were taken, of which seven were queens and 39 workers. Of the latter, 1 1 had slightly developed oocytes, but the index was 0. The tergal width and length were significantly correlated, both with each other (all N = 46; r = .375, p<.01) and with length of the discal cell (r=.567, p<.01 for tergal width; r =.726, p <.01 for length). As in the previous colony, hamu- lus number was not significantly correlated with the other mea- surements. No measurements were correlated with ovariole index (r=.026 for tergal width, —.032 for tergal length, —.219 for discal cell, and —.234 for hamulus number). Queens did not differ signifi- cantly from workers by any measure (t = 0.277 for tergal width, — 0.168 for tergal length, —1.495 for discal cell length, and —1.356 for hamulus number). When the morphometric data for each colony were pooled, signif- icant correlations were found between all measurements, but not hamulus number. When further examined, the colonies differed sig- nificantly in tergal width, with colony 2 having workers with nar- rower terga (mean = 0.73 mm versus 0.76; t = -3.353, p <.002). No correlation with ovariole index occurred in the pooled data, but all trends were negative. Summing up, it appears that queens are slightly smaller than workers but the differences are not generally significant in these young colonies. Richards and Richards (1951) reported significant differences between queens and workers in hamulus number in this species, and this result has been widely cited (e.g. Richards, 1978; Jeanne, 1980). 242 Psyche [Vol. 91 They studied 14 colonies in varying states of development, and counted hamulus number in 138 wasps drawn from an unspecified number of these colonies. Queens had a larger average number of hamuli. While their results were based on a larger sample size, these authors could not check for sperm in the spermatheca, and so classi- fied caste solely on the basis of ovariole condition. Workers were those with filamentous ovarioles and no “developed rudiments” (this term was not defined), and queens “nearly always” had some “ripe” eggs and a number of enlarged rudiments. They recognized an intermediate caste in three colonies, with “two or three rudiments more or less enlarged” These probably corresponded to our workers with some developed eggs (length>0.6 mm, index = 0). They further pointed out that their queens of one colony would have been classified as intermediates if “a typical queen” had been found. Without details on the source of their sample it is difficult to infer a complete explanation of their different results. However, their meas- ure of wing length showed the same trend as ours: queens appear to be slightly smaller. The difference in mean hamulus number, with the queens having the larger value, may be at odds with this trend (cf. Richards, 1949 on a generally positive relationship between wing length and hamulus number). Therefore, some individuals were probably misclassified. At the very least, their result does not obtain for all colonies of this species (and probably not for all of theirs), and we believe it is generally not valid. Polvbia catillifex: One colony of this species (82-02-03-1) was collected at Voltz- berg Camp on February 2. It was in primary “liana forest” on a low vine, and inconspicuous. The envelope obscured the petiole, which was attached to a twig at the base of two leaflets. These had been curled around and attached to the apices of the peripheral cells, and the envelope also was built over their bases. The total area of the comb was approximately 38 x 32 mm. The paper was brown, and composed of tough plant fragments and some granular material. It was somewhat brittle, and had been built by the edge technique. Material had been added dorsally but in an irregular fashion; the envelope was considerably thicker dorsally. None of the projections figured by Mobius (1856: pi. 8, fig. 4) and described by Richards and Richards (1951: p. 64) were present. The envelope was incom- plete ventrally, and the single comb exposed. There were approxi- 1984] Carpenter & Ross — Polistinae 243 mately 95 cells. Eggs were present in nearly all of these, attached to sides of the cells just above the base. The adults taken numbered 48, and three wasps escaped. Of the captured wasps, five were queens and 42 workers. One wasp was classified as intermediate, with an ovariole index of 1 but no devel- oped eggs. It was our impression that its oocytes were degenerate, and we included it with the workers in the statistical analyses. Mor- phometric measurements included maximum width of metasomal tergum II as well as length of the discal cell. These were found to be significantly correlated with each other (all N = 48, r = .4 1 0, p<.01), and with ovariole index (r=.375 for tergal width and .499 for discal cell length, all p<.01). Hamulus number was not corre- lated with the morphometric measures (r =.072 for tergal width and —.082 for discal cell length) or ovariole index (r=.198). The queens were found to be significantly larger than the workers in both tergal width (mean = 2.49 mm versus 2.40; t = 4.350, p<.002) and discal cell length (mean = 4.91 mm versus 4.66; t = 4.322, p<.004). They did not differ in hamulus number (t = —1 .662). Richards and Richards (1951) studied four colonies of this spe- cies. They compared eight queens and eight workers (presumably determined solely by ovarian development) for five morphometric characters, and found no differences. Width of tergum II was not among these characters. They compared average wing length and hamulus number for ten queens and 33 workers, and found no differences. Their sample sizes werev smaller than ours, and drawn from colonies in several stages of development (e.g. two colonies had two combs, and one of these had males). The fact that our colony had a larger number of workers than any of the colonies they studied, but a brood consisting entirely of eggs, suggests that this colony was the recent product of a swarm. This is supported by the incomplete envelope of our colony. Therefore the larger size of queens found by us may be characteristic of recently initiated colo- nies of this species, and not necessarily mature colonies. Braehygastra seutellaris: One colony of this species (82-02-1 1-1) was collected at Voltz- berg Camp on February 1 1. It was on a branch tip of small Clusia tree on the granite outcrop near the camp. The nest was about 1 .65 m high and readily visible. In general appearance it was of much the same form as Figs. 7 and 8 of Richards and Richards (1951). The 244 Psyche [Vol. 91 envelope was brownish-gray and quite flexible. The paper consisted of fine whitish fibers and plant epidermal fragments. Some thicken- ing of the envelope had occurred, but was not uniform. The enve- lope for each comb appeared to be the result of a single construction effort, and was attached to the edge of the combs. The entrance was at the edge of the ventral surface, and more or less continuous with the entrances to each comb. There were three combs. The oldest, basal comb contained 113 cells, of which 48 were capped and contained pupae. These were concentrated in the center of the comb. Many of the pupae were melanized, and three adults were emerging as the nest was collected. The cells around the periphery of the capped cells contained 1 1 late-instar larvae. The second comb consisted of 163 cells, of which 1 18 were capped and contained pupae and prepupae. In addition, 16 uncapped cells contained late-instar larvae. The third comb had 232 cells. There were 18 early-instar larvae scattered about the comb, and an indeterminate number of eggs at the bottom of many cells (the edges of the comb were crushed, but the cells did not appear to have eggs). The number of adults taken was 1 12; 2 escaped. Among the cap- tured wasps were 23 teneral individuals classified as ’’callow”. These were measured and dissected, but omitted from most of the statisti- cal analyses (all had filamentous ovarioles). The remainder con- sisted of 34 queens, 22 intermediates and 33 workers. Measurements included maximum width of metasomal tergum II in addition to length of the discal cell. Following Richards and Richards ( 195 1 ) for the species myersi, the number of individuals with reddening of the proximal part of the metasoma, and the degree of yellow coloring of the scutellum were recorded. Tergal width and discal cell length were significantly correlated (all N = 1 1 2, r =.27 1 , p <.0 1 ). Neither measure was correlated with hamulus number (r = — .046 for tergal width, .214 for discal cell length), although the alpha level was close to .05 for discal cell length. Discal cell length was significantly correlated with ovariole index (r=.309, p<.01), but tergal width (r=.080) and hamulus number (r =. 1 7 1 ) were not. No significant association of measure- ments with color pattern were found. When the three castes were compared by ANOVA, width of tergum II was not different (F = 1 .08, df = 88). Length of the discal cell was significantly differ- 1984] Carpenter & Ross — Polistinae 245 ent, with queens having longer wings (mean = 2.71 mm versus 2.57 for intermediates and 2.62 for workers, F = 24.54, df = 88, p <.005). Hamulus number was not different (F = 2.39, df =111; callows included), but when intermediates were lumped with workers queens were found to have a significantly higher average value (mean = 7.5 versus 7.2; t = 2.604, p<.02). When reddening of the metasoma was tested by a chi-square for differences among castes, workers were found to have a greater degree of reddening than expected, and queens were darker than expected (none had any reddening), with intermediates falling between (X2= 13.31, df=2, p<.005). Scutellar color could not be tested in this way, as almost all specimens were classified as ’’entirely yellow”; obviously there were no differences between castes. Richards (1978) reported on two colonies of B. scutellaris. He dissected 173 individuals, evidently preserved in alcohol, and com- pared 30 queens with 30 workers (drawn equally from the two nests) for forewing length, number of hamuli, number of hamuli per mm of wing length and width of tergum II. Color pattern was also compared, but he did not give details. Richards did not find differ- ences in most of these features, but detected a significant difference in hamulus number, with queens having a smaller mean number (7.07 versus 7.53). This result is directly contrary to ours, but again, there are problems with his classification into castes. He evidently separated intermediates from queens on the basis of size of the eggs (“partly developed” versus “large”), but stated that this division was “not sharp” in the first colony and termed the egg counts “rather arbitrary” in the second colony. He did not record any intermediates for the second colony but stated that some of the queens “might have been called intermediates”. It is therefore likely that some individuals were misclassified. Our sample size was larger than his, and his result of a significantly lower mean hamulus number but a slightly larger size for queens than workers (differences not signifi- cant) is contrary to the presumed general relationship (Richards, 1949), as well as to the results of our study. It is possible that the relationship varies between colonies, but there is probably a general tendency for queens to be larger in this species. Color pattern is apparently largely independent of reproductive status. Scutellar color was not associated with caste or ovariole development in our colony, and Richards and Richards (1951) found opposite tenden- 246 Psyche [Vol. 91 cies in the two colonies of the closely related species B. myersi. Their results for metasomal reddening for one nest were similar to ours (association of this and caste were not reported for the other col- ony), with reddening occurring only in workers among their dis- sected wasps. The basis of this association is unknown, but it certainly does not arise from any “effect of dissection” as the Richardses speculated, for we recorded color before dissection. Metapolybia cingulata: One colony of this species (82-02-08-a) was collected on Febru- ary 8 at Voltzberg Camp. It was oriented obliquely on a large gran- ite boulder in “mountain savannah” forest, and was about 0.5 m above the ground and well shaded. The envelope was of the same color as the lichen- and moss-grown rock face, and so the nest was very well camouflaged. It measured 6.35 by 4.45 cm in surface area, and the entrance was on the upper surface towards the lower end. The paper was green-gray, very flexible and composed of crumbly fragments in a matrix of secretion. It was uniformly thin. The secre- tion formed the “windows” (Richards and Richards, 1951) scattered throughout the envelope. The paper forming the comb was brown and composed entirely of granular material overlain by secretion. The cells were separated from the envelope, and there were a min- imum of 222 (some were destroyed during removal). No brood was found. Adults collected numbered 38, and approximately ten escaped. There were two queens, 33 workers and three males. Among the workers, four had an ovariole index ranging from 1 to 4 but none of these had any “developed” eggs (length greater than 0.6 mm). They were therefore treated as workers in the analyses. Measurements included maximum width of tergum II and length of discal cell. These were significantly correlated with each other (all n = 35, r =.586, p <.001), but not with hamulus number (r =.208 for tergal width, . 1 88 for discal cell length). None of these measures was corre- lated with ovariole index (r =. 1 79 for tergal width, .115 for discal cell length and .182 for hamulus number). Queens were not different from workers in tergal width (t =.699) or hamulus number (t= 1.2216). In length of the discal cell, the castes differed signifi- cantly by a t-test (mean = 2.74 mm for queens versus 2.82 for workers, t = — 4.776, p<.001), but not by a Mann-Whitney test (T= 14). The former test has more power, but in view of the small 1984] Carpenter & Ross — Polistinae 247 number of queens it is probably best not to attach great importance to the result. In a previous study of this species (Richards and Richards, 1951) no external indication of caste was found, and our findings are in line with this. The Larva of Brachygastra scutellaris As noted previously, Reid’s (1942) description was actually of the species myersi. A series of larvae from a second colony of B. seutel- laris was present in material from the British Museum (Natural History) examined by the senior author; the label data are: Brazil: MT, base Camp, dry forest/22. IX. 68. N. 152. OWR”. The following description is based upon both colonies. All measurements are of late-instar individuals. Length 5. 7-7. 8 mm; maximum width 2. 2-3. 5 mm. Head and body pale. Dorsal lobes (paired ridges) present on first 4 abdominal seg- ments. Pleural lobes strongly developed; no division into annulets. Integument essentially smooth. All spiracular openings subequal in size, more or less circular. Anus with ventral lips forming a bilobed slit. Head width 1.6- 1.8 mm; height (exclusive of labrum) 0.6- 0.8 mm. Head capsule very soft. Setae scattered over surface, varying in number (fig. 3). Ecdysial line and parietal bands weakly indicated, the latter smooth. Anterior tentorial pits below antennal orbits. Clypeus of the usual polistine form; frontoclypeal sulcus effaced dorsally. Labrum approximately as wide as interantennal distance, sinuate; surface roughened ventrally, with 8-10 or more papillae. Mandible as in fig. 5; 2 or 3 setae on external surface. Maxilla as in fig. 4; bases further apart than mandibular bases. Labium setate; salivary opening a raised transverse slit behind palpi. In addition to scutellaris, specimens of B. augusti, bi/ineo/ata, moebiana and lecheguana from the collection of the British Museum have been examined. The more transverse shape of the head in fig. 3 than the species figured in Reid (1942) and Dias Filho (1975) is perhaps a preservational artifact, related to the softness of the head capsule. However, it was present, with some variation, in all specimens seen. The genus is also uniform in the setae of the head capsule and the mandible with a weak ventral tooth, both diagnostic characters in Richards (1978). Presence of setae on the mandibles is 248 Psyche [Vol. 91 Figs. 3-5. Larva of Brachygasira scutellaris: 3, frontal view of head capsule; 4, mandible; 5, maxilla. another major character in Richards’ key to genera (with the excep- tion of lecheguana), but care must be used in this feature. Number of setae varied within the 2 colonies, but was always low. Conclusion Our results support the general view that caste dimorphism is poorly developed or absent among Polistinae. Where differences were found (Polybia catillifex, Brachygastra scutellaris), queens were larger than workers. Among the Polistinae where differences have been detected, queens are the larger caste, except in a few 1984] Carpenter & Ross — Po/istinae 249 swarm-founding species (Jeanne, 1980). Such morphological differ- ences between reproductives and non-reproductives may vary between colonies of the same species (Richards and Richards, 1951; Jeanne and Fagen, 1974; this paper) and within colonies over the course of a normal cycle. Moreover, although morphological caste differentiation is not pronounced in the Polistinae, behavioral and functional differentiation is well developed (West Eberhard, 1978). Lack of pronounced morphological caste dimorphism has been taken to indicate a less advanced level of eusociality in the Polistinae relative to the Vespinae (e. g. Iwata, 1976). However, by other crite- ria of social complexity (mode of suppression of nestmate reproduc- tion, colony size, duration of colonies), the swarm-founding Polistinae are at a comparable level of sociality to the Vespinae (Jeanne, 1980). The difference in degree of caste dimorphism between the two groups of highly social wasps undoubtably reflects ecological constraints of the environments in which they have evolved (Richards, 1971), with the strong queen/worker dimorph- ism of the Vespinae part of an adaptive syndrome linked with winter hibernation and a short colony life cycle (Richards, 1971). The dif- ferences in adult size between queens and workers of some polistine species suggest that in these species caste differentiation is either pre-imaginal, or that males preferentially mate with females possess- ing certain morphological attributes (or that only these females are receptive to mating). These competing hypotheses must be tested in species in which caste dimorphism has been firmly established. Although dissection and morphometric study of individuals col- lected with nests are likely to remain the primary methods of analy- sis of social structure in tropical polistines, the information provided is limited. A proper understanding of the social biology of these wasps requires in-depth behavioral studies of natural colonies. Acknowledgements Deborah R. Smith of Cornell University assisted in collection of colonies. M. C. Day of the British Museum (Natural History) arranged the loan of Brachygastra larvae. We should like to thank Janice S. Edgerly of Cornell University for the figures of the larva and G. C. Eickwort of Cornell University for critical review of the manuscript. Field work in Suriname was supported by grants from Sigma Xi, the Explorers Club and the Griswold Fund of the Department of Entomology of Cornell University. 250 Psyche [Vol. 91 Literature Cited Akre, R. 1982. Social wasps, pp. 1-105 in Hermann, H. (ed.) Social Insects IV. Academic Press, NY. Cumber, R. A. 1949. The biology of bumblebees with special reference to the production of the worker caste. Trans. R. Entomol. Soc. Lond. 100: 1-45. Dias Filho, M. M. 1975. Contribui<;ao a morfologia de larvas de vespideos sociais do Brasil (Hymenoptera, Vespidae). Rev. Bras. Ent. 19: 1-36. Jeanne, R. L. 1973. Aspects of the biology of Stelopolybia areata (Say) (Hyme- noptera: Vespidae). Biotropica 5: 183-198. 1980. Evolution of social behavior in the Vespidae. Ann. Rev. Entomol. 25: 371-396. Jeanne, R. L. and R. Fagen. 1974. Polymorphism in Stelopolybia areata (Hyme- noptera: Vespidae). Psyche 81: 155 166. MObius, K. A. 1856. Die Nester der geselligen Wespen. Hamburg. Reid, R. A. 1942. On the classification of the larvae of the Vespidae (Hymenop- tera). Trans. R. Entomol. Soc. Lond. 93: 285-331. Richards, O. W. 1949. The significance of the number of wing-hooks in bees and wasps. Proc. R. Entomol. Soc. Lond. (A)24: 75-78. 1971. The biology of the social wasps (Hymenoptera, Vespidae). Biol. Rev. 46: 483 528. 1978. The social wasps of the Americas excluding the Vespinae. Brit. Mus. (Nat. Hist.), London. Richards, O. W. and M. J. Richards. 1951. Observations of the social wasps of South America (Hymenoptera: Vespidae). Trans. R. Entomol. Soc. Lond. 102: 1-170. Velleman, P. F. and D. C. Hoaglin. 1981. Applications, Basics, and Comput- ing of Exploratory Data Analysis. Duxbury Press, Boston. West Eberhard, M. J. 1978. Temporary queens in Metapolybia wasps: nonre- productive helpers without altruism? Science 200: 441-443. THE PERSISTENCE OF ROLE IN EXTERIOR WORKERS OF THE HARVESTER ANT, POGONOMYRMEX RADIUS By Deborah M. Gordon* Department of Zoology, Duke University Durham, NC 27706 The social organization of an ant colony channels individuals into particular tasks, so that the colony continues to perform its tasks in a regular way. In other words, the colony is organized so that certain roles ( sensu Oster and Wilson 1 978), vital to its function, continue to be filled. The present study reports on role behavior in exterior workers of the harvester ant Pogonomyrmex badius. Three questions are addressed: 1) In P. badius, do particular groups of ants consistently do certain tasks? 2) Does the task per- formed by an ant depend on its age? Because P. badius workers live for about a year (Porter and Tschinkel 1982), marked individuals were observed for a longer time (up to four months) in the present study than in previous studies of Pogonomyrmex species (Holldobler 1976, Porter and Jorgensen 1981). 3) Does the task performed by an ant depend on its size? P. badius is the only polymorphic species in its genus. Since majors rarely emerged from the nest, only the role behavior of minor workers was considered in this study. However, even within the minor subcaste of P. badius, a greater size variation exists than in other Pogonomyrmex species. The relationship between size and role in P. badius minors is investigated. In the present study, the behavior of exterior workers is classified into a more detailed set of tasks than in previous studies of the genus. Foraging, i.e. food retrieval, is only one of five activities observed outside the nest. The classification of tasks used here is needed to explain other aspects of Pogonomyrmex behavior (Gordon 1983a and b). ♦Current address: MCZ Laboratories, Harvard University, Cambridge, Mass. 02138. Manuscript received by the editor April 24, 1984 251 252 Psyche [Vol. 91 Methods All experiments were conducted in the laboratory with five queen- right colonies of P. badius, each containing about 650 workers. The presence of brood inside the nest could not be ascertained. Colonies were kept in soil-filled tanks (45 X 25 X 20 cm), and fed with Bhaktar-Whitcomb diet (1970), which was placed on the soil surface in a small dish. Individual ants were marked with Testors Gloss Enamel, each with a drop on the top of the head. Single ants were kept isolated overnight after being marked, and then put back into the home colony. No adverse reactions to the marking procedure were observed. All tasks performed by ants outside the nest were classified into the following five activities (Table 1): Foraging, Nest Maintenance, Patrolling, Midden Work, and Convening. Ants inside the nest were not considered in this study. No majors were marked, since they spent almost all of their time inside the nest, probably assisting in brood care (pers. obs.). An observation record consisted of the numbers of marked and unmarked ants in each of the five categories of behavior. Observa- tions were made between 8:00 and 20:00. Each observation lasted 5-10 minutes, depending on the total number of ants outside the nest. At least one hour elapsed between successive observations. A. Continuity of role. In each of the five colonies, ants collected while performing one of the five categories of behavior were removed from the colony and marked. For example, in one colony only foragers were marked, in another only patrollers were marked, and so on. In each colony, five ants were marked every five days until 20 ants had been marked. Each group of five ants was marked with paint of a different color. Thus each colony was observed for a total of 35 days, which I call the first observation period. During this time, a total of 728 observations were made, about 145 observations on each colony. The data were analysed separately for each colony as follows. First, a two-way chi-squared procedure with four degrees of freedom (Sokal and Rohlf 1981) was used to test whether the distribution of the ratio of marked to unmarked ants depends on activity. Next, another chi-squared test, this time a two-by-two test with one degree of freedom, was performed. The latter tested whether ants were significantly likely to continue doing the activity they were doing when marked, called the “tagged activity.” Both tests used the total Table 1. Classification of activities of exterior workers of P. basius in laboratory colonies 1984] Gordon — Pogonomyrmex badius 253 00 c co JC •o a x) .E • o ■“ -o o 2 ° ^ E o no £ .S C n "O ° « O -o Ml .O C C/3 f- Ql S E 3 3 a- -c * r co 3 "2 ° 5 co -Q C/J 5 50 S .E 00 c CO >> h oo 0> G a | C .3 2 * J- 00 c u '5 1 & 3 C/3 "O C (U s g -a xj c c 3 3 O O c0 c0 00 00 C c c o CO U tc co eo {Si ^ £ £ '-5 £ £ O *' u oil - E o * 'a. 3 fc o 2 co a: c/3 U T3 . -a ^ CUM a. c. 3 3 o o 00 00 CO co £ £ 3 g ob § rrf G CT3 •— w C/3 C/5 ^ £ < ffl U 0.8993), no effect of type of activity on head width (DF = 4, SS = 0.09, F = 0.77, p > 0.5492), and a significant effect of colony of origin on head width (DF = 6, SS = 1 .30, F = 7.53, p> 0.001 ). Mean head size for all 25 workers in all five activities ranged from 1 .53 mm in one colony to 1 .78 mm in another of the seven colonies. Mean worker size increases with colony age (Oster and Wilson 1978). Differences in age among the seven colonies probably account for the effect of colony of origin on head width. Discussion At any one time, exterior workers in the P. badius colony can be divided into four groups that consistently perform certain roles for at least 35 days: 1 ) midden work and patrolling, 2) nest maintenance, 3) foraging, and 4) convening. 1984] Gordon — Pogonomvrmex bad i us 261 Table 2D. Behavior of callows while convening Activity when Marked F Subsequent Activity NM PT MW CN Marked 26 17 63 74 ** 385 Unmarked 2290 14382 4664 9731 15601 CN Dev. for Exp. -1.7 -155.2 -6.5 -43.3 + 193.8 % Dev. 6 90 12 37 101 F= Foraging; NM = Nest Maintenance; PT = Patrolling; MW = Midden Work; CN = Convening. Convening of exterior workers is observed much more often in the laboratory than in the field. Convening includes resting and mutual grooming. It probably serves a thermoregulatory function as well, because convening ants in laboratory colonies always gather in the warmest place on the terrarium surface, directly underneath a lamp. Convening has been observed in field colonies of Pogonomvrmex, but only rarely (MacKay 1981; Gordon 1983b, 1984a, b). It is possible that, in the field, convening is usually done inside the nest where conveners would be less subject to predation. The results of this study, summarized in Table 2C, suggest the diagram depicted in Figure 1. The diagram shows how role may depend on worker age in exterior workers of P. badius. It should be emphasized that the diagram is hypothetical, pending further investi- gation, and that it rests on two assumptions. In keeping with results on many other species (Wilson 1971), it is assumed that younger ants first work inside the nest, then work outside the nest for the remainder of their lives. Another assumption is that in the laboratory experiments reported on here, the death rate of ants in a particular activity depends on their age rather than on the hazards or energetic costs associated with their activity. Convening ants are clearly less active than ants in the other four activities (see Table 1). However, there is at present no empirical basis for distinguishing the other four activities in terms of the energy expended performing them. Ants doing midden work and patrolling are shown as the oldest in Figure 1 , because ants marked while doing these activities died within 262 Psyche [Vol. 91 DEATH t PATROLLING + — ►CONVENING MIDDEN WORK A FORAGING A NEST MAINTENANCE CONVENING EMERGENCE FROM NEST ECLOSION INTO ADULTS Fig. I. How role depends on age in exterior workers of P. badius. The vertical direction represents worker age. 35 days, sooner than ants in any of the other groups. It seems that convening is done at two different times in the life of P. badius individuals. One group of conveners is the youngest group shown, because callows are most likely to convene, and continue to do so for at least 35 days. The results from the second observation period suggest that convening ants become foragers, and some foragers later become conveners, as shown. Since marked midden workers did convening in the second observation period, an arrow is drawn from the patrollingand midden work group to the older convening group. There is evidence that this transition may also be made in the opposite direction: some ants marked while convening(not callows — see Part A) did midden work in the first observation period, and patrolling in 1984] Gordon — Pogonomyrmex badius 263 the second. Marked nest maintenance workers were probably in transition to patrolling during the first observation period. Later, nest maintenance ants became patrollers, as shown. The results of Porter and Jorgensen ( 198 1 ) for P. owyheei suggest a schema similar to the one of Figure 1 . Foraging, for these authors, may include midden work, patrolling, conveningand foraging as they are defined in Table 3. By examining the behavior of foragers in more detail, the present study indicates the existence of at least one group of ants older than those engaged in food retrieval: ants doing both midden work and patrolling. The relationship between these two activities will be addressed in a subsequent paper(Gordon 1984b). It would be interesting to examine role behavior in field colonies of P. badius using the classification of behavior employed in the present study. I do not attempt in this study to describe division of labor in the P. badius colony as a whole, because the behavior of interior workers is not considered. But the results raise several questions about age polyethism in exterior workers. First, there appear to be two groups of exterior workers in the colony. Ants of one group first do conven- ing and then foraging, while ants of the other first do nest mainte- nance and then midden work and patrolling. It appears that ants from either group may then become conveners. What determines to which group an ant will belong? The possibility that these groups correspond to different sizes of workers within the minor subcaste was examined, but I found no such relationship. Second, exactly how long do individual ants remain in each of their roles? For a long-lived species like P. badius, the answer to this question awaits a marking technique better than the one used in this study. After about four months, some of my marked ants were seen to have most of the paint chipped off. The unresolved questions in this study are part of a larger one: how is the distribution of workers into different roles regulated? There is considerable evidence that individuals are channelled into particular roles according to the current needs of the colony (e.g. Meudec and Lenoir 1982, Lenoir and Ataya 1983, Wilson 1983). To answer the questions raised by this study, we must come to understand the system by which the colony assesses and predicts what needs to be done. 264 Psyche [Vol. 91 Summary Exterior workers in laboratory colonies of the southern harvester ant, Pogonomyrmex haclius, were marked while performing one of the following: 1 . midden work, 2. patrolling, 3. nest maintenance, 4. convening, or 5. foraging. These activities were consistently per- formed by marked ants for at least 35 days. The results indicate that the five activities are performed by four distinct groups of ants; it appears that midden work and patrolling are done by the same ants. Three months after marking, conveners had become foragers, and nest maintenance workers had become patrollers. Younger, callow ants were most often observed convening. Midden workers and patrollers died sooner than other marked ants. These results show how role may depend on worker age. In this species the minor caste is continuously polymorphic, but no evidence for size polyethism within the minor caste was found. Acknowledgements I thank T. Meagher, M. Rausher, and especially R. Lewontin for advice on analysis of the data; S. Porter, T. Seeley, A. Lenoir, and P. Klopfer for comments on the manuscript; and J. Gregg and R. Palmer for their help throughout the project. R I I I RI NC I S Bhaktar. A. W. and W. Whitcomb. 1970. Artificial diet for rearing various species of ants. Fla. Entomol. 53(4): 229 232. Ghntry. J. B. 1974. Response to predation by colonies of the Florida harvester ant. Pogonomyrmex badius. Ecol. 55: 1328 1338. Gordon, D. M . 1983a. Dependence of necrophoric response to oleic acid on social context in the harvester ant. Pogonomyrmex badius. J. Chem. Ecol. 9(1): 105 111. 1983b. Relation of recruitment rate and activity rhythms in the harvester ant, Pogonomyrmex harbatus. J. Kans. Ent. Soc. 56(3): 277-285. 1984a. Species-specific patterns in the social behavior of harvester ant colonies (Pogonomyrmex). lnsectes Sociaux: 31(1): 74-86. 1984b. The harvester ant (Pogonomyrmex badius) midden: refuse or boundary? Ecol. Ent., 9: 403 412. HOi. i. dohi.hr, B. 1976. Recruitment behavior, home range orientation, and terri- toriality in harvester ants, Pogonomyrmex. Behav. Ecol. Sociobiol. 1: 3 44. Lknoir, A. and H. Ataya. 1983. Polythism et repartition des niveaux d’activite che/ la fourmi Lasius niger L. J. Comp. Ethol.: in press. 1984] Gordon — Pogonomvrmex badius 265 MacKay, W. 1981. A comparison of the nest phenologies of three species of Pogonomyrme.x harvester ants. Psyche 88( 1-2): 25-1 A. Mn Di e , M. and A. Lknoir. 1982. Social responses to variation in food supply and nest suitability in ants ( Tapinoma erraticuni). Anim. Behav. 30: 284 292. Ostir, G. F. and E. O. Wilson. 1978. Caste and ecology in the social insects. Princeton, NJ: Princeton University Press. Portir. S. D. and C. D. JoRCii NSHN. 1981. Foragers of the harvester ant, Pogo- nomyrme.x owyheei: a disposable caste? Behav. Ecol. Sociobiol. 9: 247-256. Porti r, S. D. and W. R. Tsohinki i . 1982. Population dynamics of harvester ant workers. In: The biology of social insects (M . D. Breed, H. E. Evans, and C. D. Michener, eds.), p. 67. Boulder, CO: Westview Press. Sokal, R. R. and F. .1. Rohi.l. 1981. Biometry. 2nd Ed. San Francisco: W. H. Freeman. Timm, N. FI. 1975. Multivariate analysis. Monterey, CA: Brooks/Cole Publishing Co. Wilson, E. O. 1953. The origin and evolution of polymorphism in ants. Qu. Rev. Biol. 28: 136 156. 1968. The ergonomics of caste in the social insects. Am. Nat. 102: 41 66. 1971. The insect societies. Cambridge, MA: Belknap Press. 1983. Caste and division of labor in leaf-cutter ants (Hymenoptera: For- micidae: Alta). III. Ergonomic resiliency in foraging by A. cephalotes. Behav. Ecol. Sociobiol.: 14( I ): 47 54. PHENOLOGY AND LIFE HISTORY OF THE FILMY DOME SPIDER (ARANEAE: LINYPHIIDAE) IN TWO LOCAL MARYLAND POPULATIONS* By David H. Wise Department of Biological Sciences University of Maryland Baltimore County (UMBC) Catonsville, MD 21228 Introduction The filmy dome spider Neriene radiata (Walckenaer) [= Linyphia ( Prolinyphia ) marginata C. L. Koch; Araneae, Linyphiidae] spins a fine, dome-shaped web in the understory vegetation of temperate forests. Early accounts suggested that this species has a typical annual life cycle; however, later research uncovered indirect evi- dence of a mixed life-history pattern in a Michigan population of N. radiata (Wise 1976). Seasonal changes in composition of this popu- lation strongly suggested that some hatchlings emerging from egg sacs laid in the spring developed rapidly, matured by August and reproduced before the end of the season. Other progeny of spring- maturing adults apparently displayed a typical annual life cycle, over-wintering as juveniles and maturing the next spring. Since this type of life cycle had not been reported often for spiders, I decided to explore it in more depth. The research reported here had two major goals: 1) To describe the phenologies of two different local popula- tions of N. radiata in Maryland, and to compare them with the phenology of the more northern Michigan population. Such a com- parison would indicate whether or not the length of the growing season might influence the species’ life history pattern. 2) To obtain direct proof of two types of juveniles in the popula- tion at the end of the spring reproductive period — rapid developers, and those that over-winter before becoming mature. The goal was to confirm the previous interpretation of the observed phenology, which was based upon indirect, nonexperimentally derived evidence. * Manuscript received by the editor August 1 , 1984 267 268 Psyche [Vol. 91 Methods The research, conducted at two sites 35 km apart, involved cen- susing undisturbed areas, manipulating the age structure of one population in a field experiment, and rearing field-collected juve- niles from both populations. Study Sites The terrain of the oak forest on the Liberty Watershed, 40 km northwest of Baltimore, Maryland, was hilly, with rocky outcrop- pings, fallen trees and small saplings offering a variety of substrates for N. radiata to attach its web. A population at the Liberty site, on a south-facing slope at 180 m elevation, was studied 1980-82. The Patuxent site was located south of Liberty on the Patuxent Wildlife Research Center near Laurel, Maryland, at an elevation of 40 m. The oak forest at Patuxent differed from that at Liberty by having several beech and some pines. The terrain was flatter, with no rocky outcroppings and with a floristically less diverse under- story. Much of the Patuxent ground cover was Vacunium sp. Rela- tively rare at Patuxent before 1980, the filmy dome spider was more common 1 98 1 -82. During these years several studies were done with the Patuxent population. Determining the Phenological Pattern in Maryland Populations During 1980-82 I monitored seasonal changes in the size-class structure of the Liberty population by repeatedly censusing 13 areas marked with short, inconspicuous stakes. At Patuxent 8 similarly marked areas were censused, but in 1982 only. Mature N. radiata were collected at Liberty (spring, 1980-81; summer, 1980) and Patuxent (spring and summer, 1981-82) by capturing every adult encountered during a search of vegetation in the vicinity of the areas set aside for censusing. These spiders provided data on yearly and seasonal changes in size at maturity and fecundity. Collected adults were anesthetized briefly with CO2, measured and then paired in jars kept in the laboratory (1980-81) or on a covered, screened porch in the forest at Patuxent (1982). The spiders were fed fruit flies ad lib. Females deposited an egg sac within one to several days after being collected. Each sac was removed, eggs were counted and the diameters of 10 eggs were measured with an ocular micrometer. In 1982 all the eggs from a sac were then placed in a 7 X 10 mm plastic vial and its open end was plugged with silk from the egg sac. The vial was placed in a humid jar, where the eggs were left 1984] Wise — Phenology of filmy dome spider 269 to develop. Many females continued to feed and deposited addi- tional sacs. However, because the frequency of multiple clutches in nature is unknown, data are given for the first sac only. Temperatures of web sites were recorded concurrently at Patux- ent and Liberty with thermocouple probes on 24 May and 8 June, 1983. Each probe was covered with black tape, so that its tempera- ture closely approximated the internal temperature of a filmy dome spider exposed to solar radiation (personal obs.). Sites selected for recording temperature originally had a N. radiata web and, as a group, spanned the range of exposure to wind and sunlight of webs at each study site. Direct Evidence for Different Rates of Development Field Experiment During the last half of the season, N. radiata populations charac- teristically contain all stages. This diversity makes it difficult to follow the growth and development of groups of juveniles. There- fore, a manipulative field experiment was designed to facilitate mon- itoring of the developmental pattern of the younger spiders in the population. Eight experimental units were placed at least 10 m apart in the undergrowth at Patuxent. Made of wood stakes that supported undulating pieces of 5.1 cm-mesh galvanized wire fencing (chicken wire), each unit was 3 m long, 1 m wide and 1 m high. Use of these standardized units made it possible to establish open, replicated populations of similar densities and microclimatic conditions, and also facilitated identification of the experimental populations. From 8-17 July 1981, filmy dome spiders were removed from the units and from surrounding vegetation. The smaller instars, all of which had emerged from egg sacs laid by spring-maturing females, were then added at random to the cleared units. On 21 July (Day 0 of the experiment) each unit contained 20-39 spiders (x = 30 ± 2), all estimated to be stages 2-4. This range represented the youngest instars in the Patuxent population on that date. Four randomly selected populations received supplemental prey from 21 July through 28 August. On 21 days each spider was given a living fruit fly; on 5 of these days each spider was given another fly during a second round of feeding. The 8 populations were censused frequently. At each census all mature spiders were removed and measured. Very small spiders that were obvious immigrants were also removed. Censusing of all units 270 Psyche [Vol. 91 continued for 17 days after supplemental feeding had ceased, and on 14 September the experiment was terminated by removing and measuring all remaining spiders. Rearing Field-Collected Juveniles A major advantage of the field experiment was that biotic and physical factors that limit growth and development were at natural levels. However, because the populations were open the develop- mental fates of individual spiders could not be monitored without error. More direct evidence of the variable developmental fates of the progeny of spring adults was obtained by collecting spiders from natural populations and rearing them individually to the end of the season or until they had matured. On 1 July 1980, 47 of the largest juveniles in the population were collected from Liberty. This collection included those offspring of spring adults most likely to complete development and reproduce that summer. In the following year a different sampling strategy was employed. In 1981, 89 of the smallest spiders (stages 1-3) in the Liberty population were collected on 6 July. These were the progeny of spring adults least likely to complete development within the season. I sampled the extemes of the size distribution in order to uncover the limits to the developmental potential of the population at a particular time. Logistical constraints prevented sampling of the entire range of size classes in a single season. In 1982 juvenile spiders were collected from Liberty and Patux- ent. As in 1981, the smallest spiders that could be found were removed, but they were collected two weeks later in the season. On 21 July I collected 41 immature spiders from Patuxent and 80 from Liberty. As in the previous two years, these spiders were also the progeny of spring adults, judging from the minimum time elapsing from maturity to hatching of progeny from the first egg sac (19-21 days; unpubl. data). In all three years the collected spiders were reared in individual containers with a super-abundance of fruit flies. Although in nature the developmental rate of some juveniles might be limited by a shortage of prey, I provided a surplus of food in order to uncover the developmental potential of each individual. In 1980-81 rearings were done in the laboratory under natural photoperiod. In 1982 the juveniles were reared on the porch at Patuxent. 1984] Wise — Phenology of filmy dome spider 271 Statistical analyses of the results of all studies were done with the UCLA BMDP programs, converted for use on Cyber computers by the Northwestern University Computing Center. Results Phenologieal Patterns in Maryland Populations Two peaks in adult abundance each season characterized the Lib- erty population (Fig. 1). Seasonal changes in the size-composition of both populations, particularly at Liberty, were similar to those found in the Michigan study (Figs. 2, 3; Wise 1976). The persistence of relatively high numbers of immature stages during the summer, and the absence of intermediate stages at the end of June, provide evidence of a polymorphic phenology: some progeny of the spring adults apparently over-wintered as juveniles, whereas others devel- oped rapidly and molted to adulthood between the end of July and the first part of September. Average adult carapace width was always significantly smaller in summer than spring (Table 1), addi- tional evidence that summer-maturing adults developed rapidly within a single season and were not the offspring of the previous summer’s adults. Females that matured in the spring laid more, but smaller, eggs than summer-maturing spiders (Table 1 ). Fecundity differences such as these could reflect different seasonally adaptive reproductive behaviors, or more simply, could have resulted entirely from repro- ductive parameters being correlated with female size. This possibility was examined. Analysis of the 1982 Patuxent data revealed statisti- cally significant correlations between number of eggs in the first sac, egg diameter and female carapace width. Stepwise multiple linear regression of the pooled seasonal data indicated that number of eggs was significantly related to both female size and mean egg size (R = .52). Most of the variation in number of eggs laid could be explained by the correlation with carapace width; addition of egg size as an independent variable increased the value of R by only .05. In a similar analysis with egg size as the dependent variable, egg number was the only statistically significant independent variable. Seasonal differences in reproductive parameters were then re- examined by comparing the adjusted means through ANCOVA of the appropriate regression equations. Differences between spring 272 Psyche [Vol. 91 • TOTAL POPULATION,* — -•' o SPIDERLINGS < 2r 1980 ION,* nm/ 0 300- 200 - -o --o' 198 1 *-^8 o, .■a^r-a-T-a- . X □ ADULT dV r-| ■ ADULT 9 ? n U Ml.-lll. APR MAY JUN JUL AUG SEPT OCT Figure 1 . Seasonal and yearly changes at the Liberty site in density of spiderlings, adults and all stages combined. Numbers on the 13 marked areas are pooled. Changes in numbers are directly related to changes in population density because these same areas were censused each date. Although censuses were not conducted as frequently in 1982, they were made often enough to reveal a pattern similar to that of previous years: scarcity of adults at the end of June and an increase by early August. and summer in number of eggs reflected seasonal differences in female size and egg size. However, the eggs laid by summer females were significantly larger than spring eggs even after correction had been made for the negative correlation with egg number (Table 2). The correlation between mean egg diameter and mean hatchling size for 46 summer females in 1982 was 0.51 (p < .001, df = 44). The structures of the Liberty and Patuxent populations differed on the same date (Fig. 3). Peaks in adult abundance occurred earlier I 980 CENSUS 198 I CENSUS NOliVindOd JO NOIldOdOdd from larger ones < 2 mm in 1981 only. Length was estimated by eye, as spiders were left in the webs. All spiders categorized as juvenile males had noticeably swollen, but incompletely developed, pedipalps. Mature spiders were recognized with practically no error. For example, from August 1979 through August 1980, 84 of 85 females found in webs off the census areas that were judged to be adults were collected and verified under a microscope as being mature. The one mistake, a penultimate female, molted to the adult stage within a few days. LIBERTY 1982 274 Psyche [Vol. 91 •— " X3 E E 3 C/T E c1- s « £ 13 J2 x 3 3 c « oo