cientist Volume 38 Summer, 1975 CONTENTS Temporal Patterns of Resource Allocation aeueeete History Phenomena ................0......06005. ance Mercedes S. Foster “The Southern Distribution of the Many-lined Salamander, Stereochilus marginatus Stephen P. Christman and Howard L. Kochman First Records of Two Percid Fishes in Florida Freshwaters........................ Ralph W. Yerger and Hal A. Beecher The Florida Spiny Lobster Fishery—A White Paper Gary L. Beardsley, T. J. Costello, Gary E. Davis, Albert C. Jones and David C. Simmons Benthic Algae of the Anclote Estuary I. Epiphytes g@iseaerass Leaves |.......:...:....:.. David Ballantine and Harold J. Humm Elemental Analysis of Selected Merritt Island Plants David H. Vickers, Roseann S. White, and I. Jack Stout Range Extensions for, and an Abnormality in, Scorpaenid Fishes Collected off the Carolinas William D. Anderson, Jr., James F. McKinney and William A. Roumillat Notes on the Introduced Gecko Hemidactylus Samenmener eel OOPEL EE BOTA 2 22 )id oooh ce acdsee sed solos he bones Robert Voss Key to the Mosses of Puerto Rico .. Harvey A. Miller and Keith W. Russell Invasion of a Renovated Pond by Walking Catfish, Clarius batrachus (Linnaeus), and Other Species .......... Lothian A. Ager Florida Junior Academy of Sciences Proceedings, 1975 Annual Meeting ....... ELSI CLES 500 I 6a a No. 3 129 140 142 QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES IN FLORIDA SCIENTIST QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Copyright © by the Florida Academy of Sciences, Inc. 1975 Editor: Harvey A. Miller Department of Biological Sciences Florida Technological University Orlando, Florida 32816 The FLoripa Scientist is published quarterly by the Florida Academy of Sciences, Inc., a non-profit scientific and educational association. Membership is open to individuals or institutions interested in supporting science in its broadest sense. Applications may be obtained from the Treasurer. Both individual and institutional members receive a subscription to the FLorma Scientist. Direct subscription is available at $10.00 per calendar year. - Original articles containing new knowledge, or new interpretation of knowledge, are welcomed in any field of Science as represented by the sections of the Academy, viz., Biological Sciences, Conservation, Earth and Planetary Sciences, Medical Sciences, Physical Sciences, Science Teaching, and Social Sciences. Also, contributions will be considered which present new applications of scientific knowledge to practical problems within fields of interest to the Academy. Articles must not duplicate in any substantial way material that is published elsewhere. Contributions from members of the Academy may be given priority. Instructions for preparation of manuscripts are inside the back cover. Officers for 1975 FLORIDA ACADEMY OF SCIENCES Founded 1936 President: Dr. W1Lu1AM H. Tarr Treasurer: Dr. ANTHONY F. WALSH Division of Research Microbiology Department University of South Florida Orange Memorial Hospital Tampa, Florida 33620 Orlando, Florida 32806 President-Elect: Dr. Patrick J. GLEASON Editor: Dr. Harvey A. MILLER 9809 W. Churchill Court Department of Biological Sciences West Palm Beach, Florida 33401 Florida Technological University Orlando, Florida 32816 Secretary: Dr. Irnvinc G. FosTER Department of Physics Program Chairman: Dr. JosEpH MULSON Eckerd College Department of Physics St. Petersburg, Florida 33733 Rollins College Winter Park, Florida 32789 Published by the Florida Academy of Sciences 810 East Rollins Street Orlando, Florida 32803 Printed by the Storter Printing Company Gainesville, Florida Florida Scientist QUARTERLY JOURNAL OF THE FLORIDA ACADEMY OF SCIENCES Harvey A. Miller, Editor Vol. 38 Summer, 1975 No. 3 Biological Sciences TEMPORAL PATTERNS OF RESOURCE ALLOCATION AND LIFE HISTORY PHENOMENA MERCEDES S. FOSTER Department of Biology, University of South Florida, Tampa, Florida 33620! Asstract: A life history consists of maintenance, growth and reproductive activities. Presumably, these activities have conflicting requirements for limited resources of time, energy and specific nu- trients. It is proposed that the amount of a resource required by an individual of a given species for any particular event probably fluctuates only within well prescribed limits. Therefore, if sufficient re- source is available over a given period of time to allow for the successful completion of two or more resource costly events, then evolutionarily, the species has determined a temporal pattern for par- titioning that resource among those events. The temporal pattern of allocation displayed should en- compass the strategy optimal for the organism in its environment. This proposal is applied to tem- poral patterns of resource allocation to the molt and breeding portions of the avian life cycle. When large numbers of avian species from several geographic regions are considered, several highly adap- tive patterns emerge. LIFE HISTORY patterns may vary considerably both within and between spe- cies (e.g., Johnson, 1963; Ricklefs, 1972). If one assumes that life history phenom- ena are subject to natural selection, then a life history should represent the op- timum strategy for the organism in its particular environment, i.e., that which will allow it to maximize its genetic contribution to subsequent generations. Gadgil and Bossert (1970) consider this question in detail. They suggest that life history phenomena belong to one of three categories: maintenance, growth, and reproduction. The former two activities are important only as they enhance re- productive success or survival for subsequent reproduction. They also sugg that because time and energy available to an organism are limited, the demz for these resources by the activities in the three categories must conflict. 7 patterns of the partitioning of time and energy, or in fact any limited reso: among the three activities are of major significance. Quantitatively, a life history pattern is definable in terms of the rela’: location of resources to its component phenomena. The significance oi quantitative considerations cannot be doubted. However, they represen’ one approach to the study of resource partitioning. Of equal or even : importance to many organisms will be the qualitative and temporal as; ‘Present Address: Museum of Vertebrate Zoology, University of California, Berkeley, Califormia 130 FLORIDA SCIENTIST [Vol. 38 resource allocation among maintenance, growth and reproduction. Qualitative analyses describe the specific types of resources allotted to each life history ac- tivity and thus could be treated as a subset of quantitative analyses. They have yet to be explored extensively. However, it is not difficult to envision the con- flicting demands for protein or other organic or inorganic nutrients of a vi- viparous parent and its unborn offspring or of the non-reproductive tissues (for growth and/or maintenance) and reproductive tissues (for growth and reproduc- tion) of an individual (Fogden, 1972; Assenmacher, 1973; Farner, 1973; Scott, 1973). I wish to deal! primarily with the temporal aspects of resource allocation— that is, when an organism uses for a specific activity those resources already al- lotted to it. I will emphasize the importance of these temporal aspects which often are neglected in the face of quantitative considerations. As indicated above, the three categories of life history activities are considered to have con- flicting demands for limited resources (Gadgil and Bossert, 1970). Evidence of this conflict consists of examples of life histories in which activities costly in terms of a particular resource are temporally separated. It is accepted generally that mutual exclusion of such activities allows for the most efficient use of the resource in question (e.g., Kendeigh, 1949; Farner, 1958). However, the temporal separation of such demanding activities only implies, but does not establish, that the activities conflict in their requirements, or if they do conflict, that this conflict is the critical factor determining their separation. The scheduling of costly events in relation to each other and to environmental events may reflect selective forces which affect characteristics contributing to fitness in other ways. Thus temporal patterns of resource allocation must be considered in terms of the total environment of the organism, including all other aspects of its life history. TEMPORAL PATTERNS OF RESOURCE ALLOCATION—One may assume that the amount of resource required by an individual of a given species for any par- ticular costly event fluctuates only within well prescribed limits. And, though circumstances may exist under which one might argue to the contrary, I would also assume that the simultaneous occurrence of two or more costly events re- quires no more resource than the sum of the amounts required for each activity alone. In other words, no net resource is expended solely as a result of the over- lap itself. If sufficient resource is available over a given period of time to allow for the successful completion of two or more costly events, then evolutionarily, the species has developed a temporal pattern for partitioning that resource among those events. Resource partitioning may be vertical (Fig. 1A) with all of the available re- source channeled into a single activity during a specific time period. Thus, the resource costly events are mutually exclusive. Alternatively, a resource may be partitioned horizontally, being divided among two or more costly events at the same time (Fig. 1B). Here the events overlap and tend to be more protracted than in vertical partitioning (Bates, 1908; Johnson, 1963; Snow and Snow, 1964). Overlap need not be complete but can exhibit any degree of intermediacy No. 3, 1975] FOSTER—PATTERNS OF RESOURCE ALLOCATION 13] (Fig. 1C-D), which may be referred to as restricted horizontal partitioning. Fi- nally, in many species, individuals will be able to overlap resource demanding events facultatively when circumstances selectively favoring their overlap arise. The temporal pattern of allocation displayed by individuals of a species will en- compass the strategy optimal for the environment in which they live. % Expended Fig. 1. Patterns of resource partitioning. Areas with vertical lines represent the portion of available resource expended on a particular event over a period of time; dotted areas represent the portion expended on another event. Total amounts expended for each activity (areas) remain constant over total time. Within total time, periods of expenditure for each event vary from pattern to pattern. A. Vertical Partitioning. Costly events are mutually exclusive. B. Horizontal Partition- ing. Costly events overlap completely. C.-D. Restricted Horizontal Partitioning. Costly events show partial overlap. 132 FLORIDA SCIENTIST [Vol. 38 To examine this proposal let us review the temporal allocation of resources to various activities in the avian life cycle. Then patterns of allocation among groups of birds living under widely differing environmental conditions can be compared. The avian life cycle is particularly suitable for consideration because extensive data are available on such cycles. It is important to remember, how- ever, that the analysis presented here with regard to temporal allocation pat- terns in avian life cycles should be similarly applicable to the life cycles of any other group of animals or plants. Birds make major resource expenditures for growth, reproduction, migration, molt and “existence activities” (modified from Kendeigh, 1949), the latter three representing various aspects of general maintenance. Because existence activi- ties are performed continuously, they will not be considered. In addition, most birds show no significant growth after reaching sexual maturity, and a large per- centage of avian species are non-migratory. Therefore, only the temporal pat- terns of resource allocation to molt and breeding, two recurring activities, are considered. Breeding generally is considered to be a particularly demanding activity for birds, though metabolic requirements for both gonadal and behavioral repro- ductive events are poorly known. Energetic and nutrient requirements for gonadal events, which have been measured for only a few birds, are summarized by Fisher (1972) and King (1972). On the basis of these studies, caloric expendi- tures appear significant only for the production of the eggs (King, 1972). This also probably is true of nutrient requirements. Unfortunately, the requirements of the behavioral aspects of reproduction “. . . cannot be satisfactorily evaluated at present’ (King, 1972). This is true of nutrient requirements as well. Studies of time budgets show, however, that time expended during reproduction to feed mates or young, to defend territories, etc., increases significantly (Verner, 1965; Verbeek, 1972). Because these activities generally require increased flight, itself a highly demanding activity (Alexander, 1968), it generally is assumed that resource requirements for these activities increase appreciably. Weight loss often is noted in individuals of various species at this point in the reproductive cycle (Davis, 1961; Helms, 1968; Fogden, 1972). Payne (1972) summarizes available data with respect to molt, which most investigators study by measuring increases in metabolic rates for molting versus non-molting birds. Results vary with species, investigator, duration of the molt, etc. Values range from 5 to 57% (Payne, 1972) with those for most passerines falling between 5 and 30%. This metabolic increase is attributable to expendi- tures for feather growth and to increased expenditures for thermoregulation (Rawles, 1960; Lustick, 1970; Payne, 1972). The nutritional requirements for re- placing feathers also have been neglected. Several studies suggest, however, that protein demands are high and amino acid requirements such that many birds must use muscle tissue as a source of these materials for feather growth (Hanson, 1962; Ward, 1969). No. 3, 1975] FOSTER—PATTERNS OF RESOURCE ALLOCATION 133 TEMPORAL PATTERNS OF MOLTING AND BREEDING—Temperate Land Birds. It is in the temperate regions of the world that the classic pattern of mutual exclusion of activities with high resource demands is most pronounced. Studies of the an- nual cycles of temperate land birds show generally that not only breeding and molt, but also migration are mutually exclusive (Tordoff and Mengel, 1956; Farner, 1958; Stresemann, 1967; Newton, 1968). The physiological mechanisms that control these recurrent events, and thus their separation, either inherently and/or in response to environmental cues, time each event so that it occurs when environmental conditions are optimal for its success and when interfer- ence from other activities is minimal. These physiological control mechanisms presumably have evolved in response to the high resource requirements of each activity and the restricted periods of high food availability (Farner, 1964). Exceptions to such patterns, exist, however. For example, in several species (Wagner, 1957; Kemper, 1959; Ligon, 1971) molt is timed to occur fairly regu- larly, while breeding is timed by an irregularly abundant resource, either food or nest materials. In addition, such species often are nomadic (ibid.) so that con- tact with appropriate conditions or available resources also is irregular. Breed- ing thus becomes more or less opportunistic, birds taking advantage of appro- priate conditions regardless of other activities (such as molt) in progress. However, in most birds, energy or other resources necessary for reproduction probably are at critical levels only for a brief part of the nesting cycle, and over- lap probably does not occur during this “resource critical” period. This also ap- pears to be true of a number of other temperate forms in which adults may continue to feed fledglings for a few weeks after the onset of the postnuptial molt (Prenn, 1937; Marler, 1956; Eaton, 1957; Dixon, 1962; Evans, 1966; Newton, 1966; Dolnik and Blyumental, 1967; Stresemann in Keast, 1968). In these forms selection seems to favor overlap to take advantage of a richer food supply. Individuals of at least two temperate species (Nucifraga columbiana, Corvus corax) may begin the annual molt before they lay their eggs (Mewaldt, 1958; Gwinner in Keast, 1968), and their molt may span much of their reproductive period. In both species large body size and long primaries dictate an early onset of molt so that it may be completed prior to a winter period of food scarcity (Stresemann, 1967). Several other species, including some birds of prey, also ex- hibit extensive overlap (Colquhoun, 1951; Watson, 1962; Raitt and Ohmart, 1966; Stresemann, 1967; Middleton, 1969; Payne, 1972). In most of these forms, molt is prolonged and coincides with a period of favorable temperatures and food supply. Overlap is largely an adaptation ensuring completion of molt prior to migration or periods of food scarcity. Tropical Land Birds. Tropical birds exist under environmental and behav- ioral regimes different from those of their temperate counterparts. For ex- ample, most species that breed in the tropics are non-migratory thus eliminating a costly activity. In addition, yearly fluctuations in many environmental fea- tures such as temperature are negligible. However, restricted periods of high food availability also seem to be the rule in tropical environments probably in re- sponse to fluctuations in rainfall. Although food levels perhaps do not fluctuate 134 FLORIDA SCIENTIST [Vol. 38 to the same degree as in temperate regions, food availability does appear to vary sufficiently to limit the breeding of tropical birds to well defined seasons (Moreau, 1936, 1950; Davis, 1953; Marchant, 1959; Miller, 1963; Fogden, 1972). As in temperate regions, molt and breeding usually are separated temporally. However, it is becoming increasingly evident that a large proportion of trop- ical birds overlap molt and breeding (Foster, 1975). Because detailed studies of the ecology and breeding biology of these species are lacking in general, it is not possible to cite specific reasons for overlap in each species. A general model pro- posed to explain this phenomenon (Foster, 1974) suggests that the enforced termi- nation of one activity (e.g., breeding) to allow for the initiation of a second (e.g., molt), vertical partitioning, would be unfavorable for most individuals in the New World and African tropics where high predation of eggs and nestlings is well documented. Under circumstances of very low nesting success the ability to re- nest several times will carry a strong selective advantage by increasing the probability of an individual successfully rearing offspring. If a bird molts and breeds simultaneously, it can substantially increase the absolute length of the reproductive period and thereby its potential number of nestings, since it can breed throughout that period designated for molt as well as during the normal breeding period. When these events overlap, molt usually is protracted (Snow and Snow, 1964). However, it need not span the entire molt-reproductive period, so the extent of overlap may vary. Birds successful early in the breeding season may show no overlap. Those particularly unsuccessful may continue to breed throughout their period of molt. Australian Arid Region Birds. The birds of this area are of particular inter- est because of the often marked irregularity of their reproductive period. Breed- ing appears to be timed primarily by rainfall which is unpredictable and gener- ally erratic (McGilp, 1923; Keast and Marshall, 1954; Immelmann, 1971; Ser- venty, 1971). Despite this, molt occurs in most species on a regular annual basis (Keast, 1968). Thus, when rain falls during the molt period of a species, the po- tential for molt-breeding overlap exists. Reports of overlap are numerous (Soderberg, 1918; Carter, 1923, 1923- 1924; Serventy and Marshall, 1957; Keast, 1959; Immelmann, 1963). Keast (1968) has reviewed in detail the timing of molt and breeding in a number of Australian dry country species. He reports many instances of overlap, though he did not find it as widespread as some earlier investigators. A number of sig- nificant features of molt among Australian birds can be recognized, however. First, molt in general tends to be protracted. Additionally, in several species, the overlap of molt and breeding results in a decrease in the rate of molt. Both would decrease the daily resource requirements for molt and minimize its interference with breeding. Finally, molt is interrupted in a few species. As Keast (1968) concludes, molt-breeding overlap is particularly advan- tageous to birds occupying a somewhat unpredictable environment, as it allows them to make maximal use of an abundant food supply. Interestingly, comparable studies of birds in other arid areas (Moreau, 1950; Marchant, 1963; Immelmann, 1967; Dawson and Bartholomew, 1968; Immel- No. 3, 1975] FOSTER—PATTERNS OF RESOURCE ALLOCATION 135 mann and Immelmann, 1968) indicate that breeding seasons, though timed by rainfall, are considerably more regular than in Australia. Shorebirds and Seabirds. The scheduling of molt and breeding in seabirds from tropical, temperate and arctic latitudes has been studied. As with other birds, breeding and molt are largely mutually exclusive (Schreiber and Ashmole, 1970; Ashmole, 1971). This separation may be accomplished in two ways. In some birds, molt and breeding periods alternate (Storer, 1960; Ashmole, 1971). In others, the molt may occupy two periods bracketing reproduction. Thus, when breeding is initiated, molt is interrupted until its completion (Stonehouse, 1962; Ashmole, 1963, 1968; Schreiber and Ashmole, 1970). Presumably this absence of overlap relates to the great amounts of energy and protein required for the production of the large eggs characteristic of seabirds (Romanoff and Romanoff, 1949; Lawrence and Schreiber, 1974), the need for adults to feed often at long distances from the nesting area, and the extremely long period of dependency of the young. As with other groups, however, exceptions have been noted. At high latitudes, where summers are short and the weather often is severe and unpredictable, molt and breeding overlap in several species (Maher, 1962; Warham, 1962; Holmes, 1966; Stresemann, 1967; Ingolfsson, 1970; Ashmole, 1971). Overlap may be broad (Johnston, 1961), or the bulk of feather replacement may follow the main breeding effort (Holmes, 1971). Some species thus are able to complete two resource costly events during the short period of maximum food production. Sim- ilar adaptations are found among temperate forms (Murphy, 1936; Johnston, 1956; Payne, 1965). Tropical species which exhibit overlap do so irregularly; molt and breeding are timed so that they usually are independent. However, if condi- tions that trigger breeding arise while the molt is in progress, both activities may proceed simultaneously (Ashmole, 1962; Schreiber and Ashmole, 1970). Discussion—When large numbers of avian species are examined, several seasonal patterns of resource allocation between molting and breeding emerge. All appear adaptive for the birds in the areas where they occur. The best known and probably most common pattern is vertical resource partitioning. Here events requiring large expenditures of resources are temporally separated to minimize physiological strain and to insure most efficient use of productive energy. This pattern is found among all types of birds from all geographic areas. In most temperate land birds the separation of breeding and molt probably is re- inforced by the occurrence of a long distance migration. Breeding terminates prior to molt, which in most forms is completed prior to migration. Thus, a min- imum of several months is available even to young produced late in the season to grow and mature prior to the time when they must migrate. If molt-breeding overlap were to extend the breeding season, late hatching young would have less time to develop before migration or the onset of winter and would be more likely to perish (Goddard and Board, 1967; Kluyver, 1971). An analagous situa- tion apparently occurs in some Southeast Asian forms in which migration is “re- placed” by a severe period of food scarcity (Fogden, 1972). 136 FLORIDA SCIENTIST [Vol. 38 A second pattern is found in most of the temperate species that show overlap. Many utilize an irregularly abundant food source, and breeding occurs irregu- larly in response to its appearance. Molt, however, occurs on a regular schedule. If conditions favorable for breeding arise while the birds are molting, they still are capable of breeding. This pattern is found also in many Australian desert birds and some seabirds. Species with very low nest success due to high predation or other causes, par- ticularly those of the African or New World tropics, commonly exhibit the third pattern. Here, molt-breeding overlap serves to extend the potential reproduc- tive period by allowing breeding to continue through the period devoted to molt. Thus, when necessary, repeated renesting is possible. In both the second and third patterns, overlap is facultative. In species that respond to irregularly occurring conditions favorable for breeding, essentially all or none of the individuals will exhibit overlap in any given breeding season. Whether or not an individual of a species having the third pattern exhibits over- lap will depend upon his own degree of nesting success. Thus the number of in- dividuals showing overlap in any given season will vary. In both these instances, resource partitioning may be horizontal or restricted horizontal. The fourth pattern is found in species whose food supply is extremely abun- dant, but only over a period of limited duration. In these forms, molt may en- croach on breeding to a varying degree so that both events may be completed within the favorable season. This encroachment may be manifest by a shortening of the reproductive period or by an overlap of molt and breeding. The pattern is particularly common among high latitude shorebirds and seabirds, though it also occurs in some temperate land and sea forms. Resource partitioning may be horizontal, but usually is restricted horizontal. ConcLusions—It often is assumed that in all organisms selection favors the separation of events with large demands for resources as a physiological adapta- tion. At the same time, however, selection is acting to better adapt the organism to the particular environment in which it lives. The direction in which selection moves will depend upon the ecological characteristics of the area. Some selective forces will operate to reinforce the separation, that is the vertical distribution of resources among costly events. Others will act antagonistically, in favor of horizontal resource partitioning and overlap. The resulting temporal distribution will represent a compromise between the opposing selective pressures. Thus it is not surprising to find several temporal patterns of resource allocation. Tem- poral as well as qualitative and quantitative aspects of resource partitioning are adaptive and will contribute to the fitness of an organism in a particular environ- ment. ACKNOWLEDGMENTS—I wish to thank Marvin R. Alvarez, Frank E. Friedl, Charles E. King, Roy W. McDiarmid, Andrew J. Meyerriecks, Gerald G. Robin- son and Glen E. Woolfenden who read the manuscript and offered critical com- ments and suggestions. I also am indebted to Ing. Eladio Carmona B., Ing. Mauro Molina U., and Carlos Gutiérrez B. of the Costa Rican Ministry of Agriculture and Jorge Campabadal of the Organization for Tropical Studies for other assist- No. 3, 1975] FOSTER—PATTERNS OF RESOURCE ALLOCATION 137 ance. Andrew D. Shumaker kindly aided in the preparation of the figure. Por- tions of this work were completed during the tenure of a National Science Foun- dation Predoctoral Fellowship and a grant from the Frank M. Chapman Me- morial Fund of the American Museum of Natural History. LITERATURE CITED ALEXANDER, R. M. 1968. Animal Mechanics. Sidgwick and Jackson. London. AsHMOLE, N. P. 1962. The black noddy Anous tenuirostris on Ascension Island Pt. 1. General biol- ogy. 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London 155:185-200. Keast, J. A., AND A. J. MaRSHALL. 1954. The influence of drought and rainfall on reproduction in Australian desert birds. Proc. Zool. Soc. London 124:493-499. Kemper, T. 1959. Notes on the breeding cycle of the red crossbill (Loxia curvirostra) in Montana. Auk 76:181-189. KENpEIGH, S. C. 1949. Effect of temperature and season on energy resources of the English sparrow. Auk 66: 113-127. Kine, J. R. 1972. Energetics of reproduction in birds. Pp. 78-107. In Farner, D. S. (ed.). Breeding Biology of Birds, Natl. Acad. Sci. Washington, D. C. Kuiuyver, H. N. 1971. Regulation of numbers in populations of great tits (Parus m. major). Proc. Adv. Study Inst. Dynamics Numbers Pop. (Oosterbeek, 1970), pp. 507-523. LawreENceE, J. M., AND R. W. ScurREIBER. 1974. Organic material and calories in the egg of the brown pelican, Pelecanus occidentalis. Comp. Biochem. Physiol. 47A:435-440. Licon, J. D. 1971. Late summer-autumnal breeding of the pinon jay in New Mexico. Condor 73: 147-153. Lustick, S. 1970. Energy requirements of molt in cowbirds. Auk 87:742-746. Mauer, W. J. 1962. Breeding biology of the snow petrel near Cape Hallett, Antarctica. Condor 64:488-499. Marcuanr, S. 1959. The breeding season in S. W. Ecuador. Ibis 101:137-152. _________.. 1963. The breeding of some Iraqui birds. Ibis 105:516-557. Maker, P. 1956. Behaviour of the chaffinch Fringilla coelebs. Behav., Suppl. 5:147-150. McGI~p, J. N. 1923. Birds of Lake Frome district, South Australia. Emu 22:237-243, 274-287. Mewa pt, L. R. 1958. Pterylography and natural and experimentally induced molt in Clark’s nut- cracker. Condor 60: 165-187. MippteTon, A. L. A. 1969. The moult of the European goldfinch near Melbourne Victoria. Emu 69:145-154. Mitter, A. H. 1963. Seasonal activity and ecology of the avifauna of an American equatorial cloud forest. Univ. California Publ. Zool. 66:1-78. Moreau, R. E. 1936. Breeding seasons of birds in East African evergreen forest. Proc. Zool. Soc. London 1936:631-653. ________. 1950. The breeding seasons of African birds—1. land birds. Ibis 92:223-267. Murphy, R. C. 1936. Oceanic birds of South America, vol. II. Macmillan Co. New York. NewrTon, I. 1966. The moult of the bullfinch Pyrrhula pyrrhula. Ibis 108:41-67. . 1968. The temperatures, weights, and body composition of molting bullfinches. Condor 70:323-332. Payne, R. 1965. The molt of breeding Cassin auklets. Condor 67:220-228. No. 3, 1975] FOSTER—PATTERNS OF RESOURCE ALLOCATION 139 . 1972. Mechanisms and control of molt. Pp. 104-155. In Farner, D. S., J. R. Kinc, anp K. C. Parkes (eds.). Avian Biology 2. Academic Press. New York. PRENN, F’. 1937. Beobachtungen zur Lebensweise der Felsenschwalbe (Riparia rupestris). J. Ommithol. 85:577-586. Rarrt, R. J., AND R. D. Onmart. 1966. Annual cycle of reproduction and molt in Gambel quail of the Rio Grande valley, southern New Mexico. Condor 68:541-561. Raw es, M. E. 1960. The integumentary system. Pp. 190-240. In Marsna tL, A. J. (ed.). Biology and Comparative Physiology of Birds 1. Academic Press. New York. Ricxters, R. E. 1972. Latitudinal variation in breeding productivity of the rough-winged swallow. Auk 89:826-836. Romanorr, A. L., anp A. J. Romanorr. 1949. The Avian Egg. John Wiley. New York. SCHREIBER, R. W., anp N. P. AsHMOLE. 1970. Sea-bird breeding seasons on Christmas Island, Pa- cific Ocean. Ibis 112:363-394. . SERVENTY, D. L. 1971. Biology of desert birds. Pp. 287-339. In Farner, D. S., J. R. Kinc, anp K. C. Parkes (eds.). Avian Biology 1. Academic Press. New York. AND A. J. MarsHALL. 1957. Breeding periodicity in western Australian birds: with an account of unseasonal nestings in 1953 and 1955. Emu 57:99-126. Scott, M. L. 1973. Nutrition in reproduction—direct effects and predictive functions. Pp. 46-68, 73-74. In Farner, D. S. (ed.). Breeding Biology of Birds. Natl. Acad. Sci. Washington, D.C. Snow, D. W., anv B. K. Snow. 1964. Breeding seasons and annual cycles of Trinidad land-birds. Zoologica 49: 1-39. SODERBERG, R. 1918. Results of Dr. E. Mjoberg’s Swedish scientific expeditions to Australia 1910- 1913. XVIII. Studies of the birds in north west Australia. Kungl. Svenska Vetenskapsakad. Hadl. 52:1-116. STONEHOUSE, B. 1962. The tropic birds (genus Phaethon) on Ascension Island. Ibis 103b:124-161. Storer, R. W. 1960. Evolution in the diving birds. Proc. XII Internat]. Omithol. Cong. pp. 694-707. STRESEMANN, E. 1967. Inheritance and adaptation in moult. Prox. XIV Internatl. Ornithol. Cong. pp. 75-80. Torvorr, H. B., anp W. R. Dawson. 1965. The influence of daylength on reproductive timing in the red crossbill. Condor 67:416-422. AND R. M. MENGEL. 1956. Studies of birds killed in nocturnal migration. Univ. Kansas Publ. Mus. Nat. Hist. 10: 1-44. VeERBEEK, N. A. M. 1972. Daily and annual time budget of the yellow-billed magpie. Auk 89:567-582. VERNER, J. 1965. Time budget of the male long-billed marsh wren during the breeding season. Condor 67:125-139. Wacner, H. O. 1957. The molting periods of Mexican hummingbirds. Auk 74:251-257. WaruaM, J. 1962. The biology of the giant petrel Macronectes giganteus. Auk 79:139-160. Watson, G. E. 1962. Molt, age determination, and annual cycle in the Cuban bobwhite. Wilson Bull. 74:28-42. Florida Sci. 38(3):129-139. 1975. Biological Sciences THE SOUTHERN DISTRIBUTION OF THE MANY-LINED SALAMANDER, STEREOCHILUS MARGINATUS STEVEN P. CHRISTMAN AND HOWARD I. KOCHMAN Department of Zoology, University of Florida, Gainesville, Florida 32611 Apstract: The species is reported for the first time from Florida and additional stations are cited for Georgia. Tue plethodontid salamander, Stereochilus marginatus (Hallowell), has been reported from bald cypress and gum swamps along the Atlantic Coastal Plain from southern Virginia to Georgia (Rabb, 1966). Stereochilus is a small, secretive salamander, seldom observed, and in general collected only with some diffi- culty. Consequently, the limits of its geographic distribution in the southern United States are poorly known, and its occurrence in Georgia (including the type-locality in Liberty County) has been questioned (Neill, 1957; Conant, 1958; Rabb, 1966). Recently, Wharton et al. (1973) have reported the discovery of Stereochilus from southeastern Georgia near the upper reaches of the Okefe- nokee Swamp. They cite their southerly range extension as the first record of the species from the Gulf of Mexico drainage. Recent collecting in Georgia by Gerald Williamson and members of the Sa- vannah Science Museum Herpetology Club, and by D. Bruce Means and the writers has established the fact that S. marginatus is widespread throughout the Georgia Coastal Plain. In addition, we have now collected it from two lo- calities in northern Florida, representing the first records of the species from that state. Stereochilus marginatus is now known from Baker County, Florida, and the following counties in Georgia: Effingham, Chatham, Bryan, Liberty, Long, Wayne, Glynn and Ware. Its distribution is probably continuous throughout the Atlantic Coastal Plain from South Carolina to northern Florida. Figure 1 shows the known localities of Stereochilus marginatus in Georgia and Florida. The Florida localities reported herein are drained by a small tributary of the St. Mary’s River, and are thus a part of the Atlantic drainage. Whether or not the Ware County, Georgia, locality reported by Wharton et al. (1973) is a part of the Gulf drainage is probably moot. The upper Okefenokee is a flat, low-lying swamp, and no doubt the waters of the Satilla (Atlantic drainage) and the waters of the Suwannee (Gulf drainage) intermingle during periods of high water. Stereo- chilus has not, however, been found in apparently suitable habitat of the Suwannee River drainage, just 10 miles south of the Baker County localities. Nine specimens of S. marginatus from Baker County (UF 32561-32569) and one from Ware County (UF 32560) have been deposited in the Florida State No. 3, 1975] CHRISTMAN AND KOCHMAN—MANY-LINED SALAMANDER 14] Pa (Nee ir 312 29° ocean KILOMETERS y >: TAN BiG eulihio 100)» 290 yf hs Sy ase| 83°| far. ul oh Fig. 1. Distribution of the many-lined salamander, Stereochilus marginatus, in Georgia and Florida. Museum. Seven specimens from Liberty County are in the personal collection of D. Bruce Means, and 179 specimens from seven counties in Georgia are housed at the Savannah Science Museum. ACKNOWLEDGMENTS—We thank Gerald Williamson of the Savannah Science Museum for supplying us with most of the locality records of Stereochilus in Georgia. LITERATURE CITED Conant, R. 1958. A Field Guide to Reptiles and Amphibians of the United States and Canada East of the 100th Meridian. Houghton Mifflin Co., Boston. NeiLt, W. T. 1957. Distributional notes on Georgia amphibians and some corrections. Copeia 1957(1):43-47. Rass, G. B. 1966. Stereochilus and S. marginatus. Cat. Amer. Amphib. Rept. p: 25.1-25.2. Wuarton, C. H., T. FRENcu, anp C. RuckDESCHEL. 1973. Recent range extensions for Georgia am- phibians and reptiles. HISS News J. 1(1):22. Florida Sci. 38(3):139-141. 1975. Biological Sciences FIRST RECORDS OF TWO PERCID FISHES IN FLORIDA FRESHWATERS RALPH W. YERGER AND HAL A. BEECHER Department of Biological Science, Florida State University, Tallahassee, Florida 32306 Asstract: A male Stizostedion canadense was taken from the Apalachicola River and a male Ammocrypta asprella was collected from the Escambia River. THE expanding list of fishes known from the freshwaters of Florida was discussed by Yerger and Suttkus (1962). They noted that virtually all of the re- cent additions were collected in the Panhandle region of western Florida, and predicted that future surveys of the larger rivers would likely disclose addi- tional unrecorded species. The state list has been augmented greatly in the past decade by the introduction of more than three dozen species of exotic, tropical fishes into the southern half of the peninsula (Courtenay et al, 1974). We now report the occurrence of two additional species of the perch family (Percidae) in north-western Florida, one an introduction, the other an apparently indige- nous form. 1. Stizostedion canadense (Smith)—Sauger. A male, 241 mm standard length, was caught on hook and line in the Apalachicola River at Chattahoochee, Gadsden County, on February 3, 1962, by E. A. Burkett while fishing from the catwalk just below the Jim Woodruff Dam. This fish was identified by John T. Brown and W. Keith Byrd, former employees of the Florida Game and Fresh Water Fish Commission, and later donated to the Florida State University Fish Collection (FSU Catalog no. 7592). The sauger is not native to Georgia (Dahl- berg and Scott, 1971) or to Florida. The species was introduced into Bartlett’s Ferry and Oliver reservoirs (Chattahoochee River) near Columbus, Georgia, by the Georgia Game and Fish Division in January 1961 (John T. Brown and Don Johnson, personal communications). The Florida fish undoubtedly represents an individual which made its way downstream approximately 125 miles from the site of introduction. To the best of our knowledge, no other saugers have been taken in Florida waters. Several have been recovered from Lake Seminole, the reservoir formed by the Jim Woodruff Dam, but the sauger stockings in the Chattahoochee River of Georgia are considered to be unsuccessful (Don Johnson, personal communication). 2. Ammocrypta asprella (Jordan)—Crystal darter. The range of the crystal darter extends from southern Minnesota and Wisconsin to Ohio and south to Tennessee, Arkansas, Oklahoma, Louisiana and Mississippi (Moore, 1968), and to several rivers in the Mobile Bay drainage of Alabama (Smith-Vaniz, 1968). Dur- ing an ecological survey of the Escambia River, a subadult male 63 mm standard length (FSU No. 21354), was collected during the night of April 7-8, 1972 by No. 3, 1975] YERGER AND BEECHER—PERCID FISHES 143 W. C. Hixson, Charles A. Lowery, and Gilbert McGhee using a boat-mounted electric shocker. The collection site was the main channel of the Escambia River, approximately 1.6 km downstream from the bridge on State Highway 4, about 2.7 km east of Century, Escambia County. Although the precise col- lection site could not be determined because of the nature of the night oper- ation, the specimen was taken from waters 2-5 m deep, and over a clay-mud to sand bottom with some gravel. Other darters collected in this general locality were Ammocrypta beani, Percina caprodes, P. nigrofasciata, P. uranidea, Etheo- stoma davisoni, E. swaini, E. histrio, and Etheostoma species (orangeside darter). The 13 month survey included 75 hr of electrofishing and numerous seine hauls. The locality was revisited in the spring of 1973 but attempts to seine additional specimens were fruitless. The crystal darter was not listed by Bailey, Winn, and Smith (1954) in their report on the fishes of the Escambia River. The collection of a single specimen raises the question whether this species is native to the system or whether it was introduced. We consider its introduction improbable because the species is not a common baitfish, food fish, or aquarium species. More likely it is indigenous, but because it inhabits large flowing streams and frequents strong currents (Smith-Vaniz, 1968), and because its population density is probably low at the periphery of its range, collection of the crystal darter would be an uncommon event in this drainage system. ADDENDUM—Since the submission of the original manuscript, two additional adult specimens of Ammocrypta asprella were collected from the Escambia River on 28 November 1974 by Hal Beecher, John Stowe, and Dave Buecker. Both specimens (FSU 23033 and 23345) were seined on slip banks at consecutive sharp bends in the river, in the same vicinity as the first specimen reported in the paper. The bottom consisted of many pebbles (1-2 cm in diameter) over fine clay-sand. Fishes collected in the same seine haul with FSU 23033 included Car- piodes velifer, Notropis texanus, N. venustus, N. longirostris, Hybopsis amblops, Ericymba buccata, Ammocrypta beani, and Etheostoma species (orangeside darter). LITERATURE CITED Bartey, R. M., H. E. Winn, ann C. L. Smitu. 1954. Fishes from the Escambia River, Alabama and Florida, with ecologic and taxonomic notes. Proc. Acad. Nat. Sci. Phila. 106:109-164. Courtenay, W. R., Jr., H. F. San~man, W. W. Mitey II, anp D. J. Herrema. 1974. Exotic fishes in fresh and brackish waters of Florida. Biol. Conservation 6:292-302. Dauperc, M. D., anv D. C. Scott. 1971. The freshwater fishes of Georgia. Bull. Georgia Acad. Sci. 29:1-64. Moores, G. A. 1968. Fishes. In: Buarr, W. F., A. P. Biarr, P. BRopkors, F. R. CAacLe, AND G. A. Moore. Vertebrates of the United States. 2nd ed. McGraw-Hill. New York. SmitH-Vaniz, W. F. 1968. Freshwater Fishes of Alabama. Auburn Univ. Agr. Exp. Sta. Auburn, Ala. vii+211 p. Yercer, R. W., anv R. D. Sutrkus. 1962. Records of freshwater fishes in Florida. Tulane Stud. Zool. 9:323-330. Florida Sci. 38(3):142-143. 1975. Conservation THE FLORIDA SPINY LOBSTER FISHERY A WHITE PAPER! Gary L. BEarpsLey, T. J. CosTELLo, Gary E. Davis, ALBERT C. JONES, AND Davip C. SIMMONS American Institute of Fishery Research Biologists, 75 Virginia Beach Drive, Miami, Florida 33149 Asstract: The Florida spiny lobster fishery is faced with declining catch rates resulting from increasing fishing pressure by commercial and recreational fishermen. Conflicts between users have developed and economic returns, at least to the commercial fishery, are not optimal. Management action is suggested. Phase 1 of a management program should allocate the resource in an effective manner between recreational and commercial interests, adopt uniform interstate regulations to pro- tect the resource, and augment the collection of fishery statistics for both recreational and commer- cial harvests. Phase 2 of the program should establish a management scheme to obtain the optimum sustainable yield. . SPINY LOBSTERS, Panulirus argus, presently attract thousands of recreational divers to the Florida Keys and provide Florida’s commercial fishermen with their second most valuable catch, valued at about $15 million in 1974. Both the sport and commercial lobster fisheries were established in southern Florida by 1920. It was not until the late 1940’s that the commercial trap fishery began to grow rapidly. Increases in the recreational harvest began in the late 1950's, with the explosive growth of skin diving in that period. These two interests now actively compete for the limited number of lobsters available in Florida waters. As a re- sult of the enormous growth of these two groups and their respective harvests, a decrease in the lobster stocks has become evident in recent years. New measures are required if we expect to harvest a sustained yield from the available stocks, while at the same time perpetuating the lobster resource and maximizing its con- tribution to Florida’s economy. Florida’s legal regulations involving gear restrictions, seasons of capture, and condition and size of animals pertain to only one species of spiny lobster, Panu- lirus argus, which is distributed from Bermuda and North Carolina to Brazil. Other species of spiny lobster are present in Florida’s marine environment, but form only a minor proportion of the total lobster catch. THE PROBLEMS 1. Declining Abundance. What data are available indicate that there have been serious declines in the catch per unit of fishing effort in recent years (Sea- man and Aska, 1974). Divers find that areas which previously had good lobster 'This paper has been approved by the membership of the Florida District of the American Institute of Fishery Research Biologists, and constitutes a position paper from that organization. No. 3, 1975] ‘BEARDSLEY ET AL.—SPINY LOBSTER FISHERY 145 fishing now have few lobsters. Commercial fishermen who in early years oper- ated only 200 traps now use as many as 2000 traps to make the same harvest. In our opinion, based on communication with both fishermen and scientists, this intensified fishing pressure creates an exploitation rate for the Florida popula- tion that is high enough so that most lobsters at or near legal size in the Florida Keys are being caught each year. 2. Low Economic Return. One cause of the decline in catch rates has been the continuing increase in effort in the commercial and recreational fisheries. Unregulated entry has allowed addition of more and more gear to a fishery which is already harvesting near maximum yields. More gear brings higher costs and, with a less than equivalent increase in overall harvests, lower net financial returns to individual fishermen. State of Florida records show an increase of more than 1000% in the number of lobster traps fishing in the 1969-70 season as com- pared to the number of traps fishing in the 1955-56 season. For the same period total catch increased only 100%. Present earnings and net financial returns of commercial vessels are not high (Noetzel and Wojnowski, 1975; Dept. Food and Resource Economics, Univ. Florida, 1975, unpublished data). Under existing conditions, neither the commercial nor the recreational fishery can be expected to meet its full economic potential. Commercial fishermen are being forced out of the fishery because of low return on their investment, and income derived from recreational diving in south Florida is in danger of being reduced. The rapidly intensifying conflict between recreational and commercial in- terests in the fishery must be resolved so the lobster resource may provide max- imum benefit to society. If these problems are not addressed soon, not only may the ability of the fishermen to economically harvest this high quality protein source for society be eliminated, but the reproductive potential of the population _ may be reduced. 3. Ineffective Regulations. The present management scheme of Florida prohibits taking gravid females, all animals with carapace lengths less than 76 mm (3 inches), and all animals during the peak breeding season, April through July; and also restricts the type of fishing gear used. These regulations have the primary purpose of protecting the lobster stocks as biological entities, but do little or nothing to improve the economics of the industry. Furthermore, enforce- ment of these few rules is difficult because of the large area involved and the high potential gain/penalty ratio for the offender. Trap theft is a major problem. The success of the management scheme is also limited by the political boundaries of Florida’s territorial waters, which have no rational relationship with the nat- ural distribution of P. argus. Management is ineffectual in controlling the local high seas harvest or the international recruitment potentials. 4. Inadequate Fishery Statistics. Any management scheme, other than com- plete laissez faire, requires detailed, accurate record keeping of both recrea- tional and commercial harvests. Recreational harvest statistics do not exist. Pres- ent commercial harvest figures alone are inadequate to manage the fishery. Measures of fishing effort, the age and size structure of the population, detailed geographical and temporal information on the harvest, and an economic pro- file of the industry are required to monitor the fishery. 146 FLORIDA SCIENTIST [Vol. 38 5. Lack of Natural History Information. Basic biological information is not available to manage the fishery for optimum sustainable yield (OSY), defined as the largest net economic return consistent with the biological capabilities of the stock, as determined on the basis of all relevant economic, biological, and en- vironmental factors (Roedel, 1975). Until the population dynamics, growth rates in the wild, migratory patterns, age at maturity, and fecundity are better under- stood, no reliable estimate of the crop of lobsters that may be optimally harvested will be possible. Also, many facets of the life history of P. argus are not known with the accuracy and precision needed to recommend specific management strategies. For example, is enhancement of postlarval survival by the use of arti- ficial habitats feasible? What is the biological capacity of various habitats for both adult and juvenile lobsters?P—and how can this information be applied to ob- tain the optimal use of the resource for both recreational and commercial inter- ests? Lobsters which eventually grow up and are harvested in the Florida fishery may have originated from outside Florida waters. There is good reason to be- lieve that during the several months that young lobsters live in the surface waters of the open ocean, many of them drift hundreds or thousands of miles away from where they were hatched. If Florida’s fishery depends on recruitment from distant areas, it is important that other states and nations, as well as Flor- ida, take steps to insure that their lobster populations are managed in such a way that successful reproduction is assured. Resolution of this question of source of recruits will require scientific investigations of the identity of larvae of the palinurid family and of their behavior. MANAGEMENT ACTION NEEDED The management action program recommended here is two phased. Phase 1 can be instituted at the present time, and consists of (1) the allocation of the re- source in an effective manner, (2) the adoption of uniform regulations, and (3) the collection of necessary fishery statistics. Phase 1 has as its goal the establishment of management control over the fishery. Phase 2 is a subsequent phase of the management action program which can be initiated after certain biological and economic information requirements are met. Phase 2 has as its goal the establish- ment of an optimum sustainable yield fishery. PuasE 1. 1. Allocate the Resource. Immediate consideration should be given for some form of allocation of the Florida lobster resource in order to in- crease the economic profitability of the fishery. A moratorium on commercial fishing permits might be a first step in stabilizing the number of units of gear in the commercial fishery at a level consistent with good fisheries management and fair dollar return to the fishermen. Allocation also is a means of optimizing the return from the recreational fishery. Management by limited entry operates suc- cessfully in the lobster fisheries of western Australia (Anonymous, 1974b; Bowen, 1971). It has recently been introduced or proposed for the salmon fisheries of Alaska and British Columbia and the lobster fisheries of Maine, the maritime provinces of Canada, and Turks and Caicos (Campbell, 1973; Dow, et al., 1975; No. 3, 1975] BEARDSLEY ET AL.—SPINY LOBSTER FISHERY 147 State of Alaska, 1974; R. Stevens, personal communication, May 22, 1973). There is ample legal precedent for basing management programs on the economic well- being of the industry as well as on protection of the resource. The greatest bene- fits of management are in cost reduction to the industry rather than through in- creased production from the resource base (Herrington, 1972). In Florida, or- ganized groups of commercial fishermen have recently expressed a desire to adopt limited entry schemes for lobster (Seaman and Aska, 1974). These fisher- men are acutely aware of, and suffer from, the economic effects of declining catch rates and the associated increased costs and effort necessary to maintain the present level of catch. Inherent in any allocation scheme must be a partitioning of the resource be- tween recreational and commercial interests. The competing interests of com- mercial trap fishermen and recreational divers must be resolved. The lobster fishery is a livelihood to some. To others its recreational use represents a needed respite from their daily routines and a valuable addition to the local economy. These uses need to be placed in perspective and potential conflicts resolved. Allocation will require certain socio-economic decisions concerning who should benefit from publicly owned resources. Once these decisions are made and implemented, the remaining management decisions are purely technical ones. These technical decisions concern mainly the legal regulations and the management data necessary to operate the system. We believe that these tech- nical decisions can be made from information which is already available or which can be acquired at reasonable cost and effort. Thus, if an allocation scheme is agreed upon and adopted, management can be implemented immediately. 2. Adopt Uniform Regulations. The international nature of this fishery must be recognized in all attempts at management, and uniformity of regulations es- tablished between all states and nations involved in harvest of spiny lobsters. Fortunately, several Caribbean countries, including the Bahamas, have modeled their conservation laws after those of Florida. While general uniformity of regu- lations is desirable, it is recognized that specific details may vary from area to area. In 1975 a proposal was submitted to the United States Congress (94th Con- gress, lst Session, H.R. 2473) to provide uniform interstate regulations to protect juvenile and egg-bearing spiny lobsters. Uniform laws are an important first step towards providing an adequate basis for protection of the biological resource. 3. Collect Detailed Fishery Statistics. Detailed fishery statistics must be gathered for both sport and commercial harvests. These should include: (1) the quantity and value of the spiny lobster catch by trip, area of capture, and method of capture; (2) a measure of fishing effort for each trip; (3) length or weight fre- quency description of the catch; (4) vessel identification/description informa- tion; (5) operating costs and net economic return for the commercial and recrea- tional fisheries. These data must be available on a timely basis (maximum 30-day delay) to effectively monitor and manage the harvest. Puase 2. Phase 2 of the management program envisions the establishment of a fishery based on optimum sustainable yield. The present situation in the lob- ster fishery, vis-a-vis management for OSY, is analogous to a city manager 148 FLORIDA SCIENTIST | [Vol. 38 having to provide services, plan for future growth, and meet fiscal responsibili- ties of any of the major cities in Florida without having adequate information on (1) the size of the population of his city, (2) the rate of movement into or out of the city, (3) the age and sex distribution of the population, (4) the socio-economic profile, or (5) the monies available from which he must budget. Phase 2 will re- quire additional research before the most appropriate management action can be initiated. Suggested topics for research are: 1. Monitor Juvenile Population. A promising approach to a realistic man- agement scheme is through monitoring of postlarval and juvenile recruitment, as is done in the Australian lobster fishery (Anonymous, 1974a; Bowen, 1971). Predictions a year or more in advance of the numbers of fishable lobsters based on postlarval or juvenile abundance are free of variations in egg production, oceanic current effects, larval mortality, or settlement habitat conditions. Since recruitment may be partially independent of local adult population densities, this approach should provide accurate predictions of available yield with little additional input. Development of an adequate postlarval or juvenile monitoring program is of high priority in the research needs of the fishery. 2. Study Natural History. Basic information is needed about the growth rates of spiny lobsters in the wild, their migratory and seasonal movements, fe- cundity, age at maturity, behavioral responses to divers, and changes in the en- vironment. These observed facts must then be integrated with a population monitoring program. Some of these data presently are being collected concur- rently with in situ surveys and scientifically managed trapping-tagging studies of the fisheries (Seaman and Jones, 1975). Other information is available only from populations unaffected by harvest pressures. For example, south Florida’s under- water parks, some of which are presently closed to lobster fishing, might be uti- lized for determination of habitat carrying capacity, impact of various manage- ment strategies on natural populations, and undisturbed rates of production. Such closed areas also protect a component of the population that will produce larvae for neighboring fisheries and adult recruitment to adjacent areas. 3. Determine Larval Abundance and Distribution. The ultimate source of recruitment to the Florida spiny lobster fishery remains an enigma. Problems of research on the pelagic larvae of spiny lobsters are formidable but solutions, al- though ultimately necessary, are not immediately imperative. Information on postlarval and juvenile recruitment is more readily available and may be suffi- cient for short-term management needs. Resolution of larval research problems and development of indices of phyllosome abundance might enable long-range forecasts of future fishery productivity to be made. Improved knowledge of phyl- losome biology would also greatly assist development of larval rearing techniques for scientific and commercial purposes. 4. Continue Cooperative Programs. The present close contact should con- tinue between State, Federal, University, and private interests involved in spiny lobster research and management. Standardization of data collection and estab- lishment of a common “data bank,” presently in progress, will provide an up- to-date, complete source of basic biological, fishery, and economic data for re- No. 3, 1975] BEARDSLEY ET AL.—SPINY LOBSTER FISHERY 149 searchers and managers. In view of the pan-Caribbean distribution of spiny lob- sters, and because of our dependency upon other Caribbean spiny lobster popula- tions for postlarval recruitment, it is imperative that good management prac- tices be fostered throughout the species’ range. CONCLUSION The spiny lobster resource is not at present providing maximum benefit to the people of the State of Florida. The fishery suffers from excess effort and low catch rates. Allocation of the resource, adoption of uniform regulations, and es- tablishment of a program to collect detailed fishery statistics are suggested as immediate actions. Management of the fishery for optimum sustainable yield should be an ultimate goal, and is suggested as a second phase of the management program. Management based on these concepts can provide substantial bene- fits, including stabilized annual production, increased economic efficiency, and reduced user conflict. ACKNOWLEDGMENTS— This White Paper was prepared by the Committee of five authors, A. C. Jones, Chairman. The authors wish to express their sincere appreciation to the many scientists and industry members who participated in the two conferences on spiny lobster held in 1974 (Seaman and Aska, 1974; Sea- man and Jones, 1975). Discussions at these conferences laid the groundwork and interest in this report. Subsequent discussions with some of the participants, es- pecially Dr. Richard Warner, added to the ideas. Nevertheless, the opinions ex- pressed in this White Paper are the sole responsibility of the authors. LITERATURE CITED Anonymous. 1974a. 1973-74 rock lobster season. Australian Fisheries 33(9):2. _______. 1974b. Fisheries Act operates from January 1975. Australian Fisheries 33(12):6-7. Bowen, B. K. 1971. Management of the western rock lobster (Panulirus longpipes cygnus George). Proc. Indo-Pacific Fish. Coun. 14(II):139-153. CAMPBELL, B. A. 1973. License limitations regulations: Canada’s experience. J. Fish. Res. Board Canada 30:2070-2076. Dow, R. L., F. W. BELL anp D. M. Harriman. 1975. Bioeconomic relationships for the Maine lob- ster fishery with consideration of alternative management schemes. NOAA Technical Report NMFS SSRF-683. pp. 44. Herrincton, W. C. 1972. Management of fishery resources for optimum returns. Would it work in the Gulf of Mexico? Proc. Gulf Caribbean Fish. Inst. Ann. Sess. 24:33-41. NoetzeEL, B. G. anp M. G. Woynowskxt. 1975. Costs and earnings in the spiny lobster fishery, Flor- ida Keys. Marine Fish. Rev. 37(4):25-31. RoepEL, P. M. (ed.) 1975. Optimum sustainable yield as a concept in fisheries management. Ameri- can Fish. Soc. Spec. Publ. 9:pp. 89. SEAMAN, W., JR. AND D. Y. Aska. (eds.) 1974. Research and information needs of the Florida spiny lobster fishery. State Univ. Syst. Florida Sea Grant Program Rept. 74-201:pp. 64. AND A. C. Jones. (eds.) 1975. Review of Florida spiny lobster research. State Univ. Syst. Florida Sea Grant Program. pp. 52. State oF ALaska, Commercial Fisheries Entry Commission. 1974. Proposed regulations, limited entry: Report to the fishermen of Alaska. July 15, 1974. pp. 26. Juneau. Florida Sci. 38(3):144-149. 1975. Biological Sciences BENTHIC ALGAE OF THE ANCLOTE ESTUARY I. EPIPHYTES OF SEAGRASS LEAVES Davip BALLANTINE AND HAROLp J. HuMM Department of Marine Science, University of Puerto Rico, Mayaguez, Puerto Rico 00708; and Department of Marine Science, University of South Florida, St. Petersburg, Florida 33701 ABSTRACT: Sixty-six species of benthic algae are recorded as epiphytes on the 4 species of sea- grasses that form extensive beds in the Anclote estuary near Tarpon Springs, Florida Gulf coast. Monthly field observations and collections were made at 6 representative stations from January to September, 1971. About 65% of all benthic algae that grow attached in the area occur as seagrass epiphytes. Ceramium byssoideum fa. alternatum is newly described. Tue broad continental shelf of the Florida Gulf coast from the Keys to Apa- lachicola supports the most extensive seagrass beds of the North American con- tinent. Since there are few rocky outcrops on the inner shelf where most sea- grasses occur, the leaves of seagrasses are the most important substrate for ben- thic algae in depths of less than 10 m (Humm 1956, 1973). As part of a general environmental research project in the Anclote River estuary at Tarpon Springs (Baird et al., 1972), a study of algal epiphytes of sea- grass leaves was carried on from January to September, 1971, a time period that included all significant seasonal changes of the flora for the year. Three seagrasses occur in abundance in the area studied and along the entire Florida Gulf coast: Thalassia testudinum Konig (turtle grass), Syringodium fili- forme Kitzing (manatee grass), and Diplanthera wrightii (Ascherson) Ascherson (shoal grass). A fourth species, Halophila engelmannii Ascherson is occasional, usually mixed with Thalassia. A fifth species, H. baillonis Ascherson, has not been recorded from the Anclote estuary but is occasional along the Florida Gulf coast, especially in deeper water (10 to 100 m). The latter two have no common name. In view of the similarity of the marine environment over the inner conti- nental shelf of the Florida Gulf coast and the Anclote River estuary, it is be- lieved that the seagrass epiphytes recorded here will include an overwhelming majority, perhaps 90%, of the species occurring in other seagrass beds between Fort Myers and Apalachicola. ENVIRONMENT—Area. If the Anclote River estuary is delimited by North An- clote Key (as the NW corner), by Bailey’s Bluff (as the NE corner), by Piney Point (as the SE corner), by the south end of Anclote Key (as the SW corner), then the total area is about 28 sq km (12.3 sq miles). This area is approximately a square of about 5.2 km (3.5 miles) about 40% of which or 11.2 sq km (4.9 sq miles), is covered by seagrasses (Zimmerman et al., 1973) as determined by aerial photog- raphy (Feigl and Pyle, 1973). Zonation. Along the mainland side of the Anclote estuary, the seagrasses ex- hibit four zones, from the beach outward, as follows: zone 1, Diplanthera, from a No. 3, 1975] BALLANTINE AND HUMM—ALGAL EPIPHYTES 151 low intertidal (spring tides) to a few cm depth, averaging about 30 m wide; zone 2, Thalassia, from about 100 to 300 mm depth, a band averaging 35 m wide; zone 3, Syringodium (dominant) with some Thalassia and Diplanthera mixed with it in a band about 0.70 to 1.5 m deep, the zone about 400 m wide; zone 4, Diplanthera, with a small amount of Syringodium, as an outer narrow band beginning about 700 m out at a depth of about 0.17 m to more than 1 m in the clearer parts of the estuary (Zimmerman et al., 1972). Physical Factors. Salinity of surface water in Anclote estuary during 1971 ranged from 26 to 32°/oc, based upon samples taken once a month. Salinities below 30 were recorded in April, September, October, and November. While sampling was too infrequent to obtain the full perturbations, the data show that Anclote estuary is similar to the adjacent Gulf of Mexico and that evaporation and tidal mixing in the area almost cancel the fresh water contribution from the Anclote River. Rainfall during 1971 was lower than 13 cm per mo in the area ex- cept for July (23), August, and September (43 each). Surface water temperatures ranged from a low of 11.2°C during February, 1971, to a high of about 32° during the summer and a decline to 17° in Novem- ber. From January through April, water temperature of the adjacent Gulf was 2-3 degrees higher; during summer and fall the difference was only about 1 degree. Tidal current velocities in the estuary ranged from about 0.15 to 0.40 m per sec at times other than slack tide. The tidal amplitude of spring tides was 0.8 m but this was often exceeded or reduced by wind direction and velocity. Light penetration in waters of the Anclote estuary was determined by means of a transmissometer employing either a 0.1 m or a 1 m light path. In general, from 45 to 60% of light striking the water surface penetrated to a depth of 0.1 m. The clearest water was usually in the northwest sector of the area (Pyle et al., 1973). Nutrients are relatively low in the Anclote estuary (Johansson and Hopkins, 1973) and are more characteristic of inshore Gulf waters than of estuaries of the Florida Gulf coast. The area is little disturbed and relatively unaffected by the town of Tarpon Springs or the metropolitan areas of Tampa and St. Petersburg. PROCEDURES—Sikx stations were established in the Anclote estuary in order to insure a sampling of all the obvious types of seagrass communities, and to insure repeated collecting in a selected area to determine seasonal changes. Collections were made at monthly intervals at each station within a 10 m radius of a buoy placed to mark the station location. Stations 1, 2, and 3 were located in the northern segment of the Anclote es- tuary from an in-shore point near the U. S. Air Force radar station and extending westward to the outer margin of seagrass stands. These three stations were lo- cated to represent the different types of seagrass communities found along this transect. Station 1 was established about 20 m from the mean low water line (west) in zone | and included the in-shore margin of the seagrass community. Diplanthera was the most abundant plant but there was some Thalassia in the outer part of 152 FLORIDA SCIENTIST [Vol. 38 the station area. Halophila was also present, but sparse. The depth was from 0.2 m to 1 m, depending upon the tidal cycle. Station 2 was established about 600 m from shore (west) where there was a dense stand of Thalassia with an admixture of Syringodium, typical of zone 2. Diplanthera and Halophila were also present in low abundance. The depth ranged, with tidal level, from 1 to 2 m. Station 3 was established near the outer margin of the seagrass community about 1000 m from the shore line (west). Diplanthera was the most abundant seagrass, but the other three were present in low abundance, typical of zone 4. Stations 4, 5, and 6 were located in the western and southern parts of the An- clote estuary on the basis of a careful reconnaissance for the purpose of locating representative types of seagrass stands differing from stations 1-3. Station 4 was established on an extensive shallow flat on the north side of the boat channel leading from the mouth of the Anclote River. Diplanthera was the dominant seagrass but there were small, scattered patches of Thalassia. The sta- tion was representative of zone 1 but differed from station 1 in its nearness to the river mouth and consequent exposure to periods of reduced salinity, stronger currents, and greater turbidity. The depth at station 4 generally ranged, with tide, from about 0.3 m to 1 m, but during January large areas of station 4 were exposed to the air during spring low tides when there was a strong NE wind. At the same time, the seagrasses were exposed to subfreezing temperatures resulting in considerable death of the erect branches and leaves of a small, pure stand of Halophila that was present near station 4. Station 5 was established about 1000 m south of station 4 off the SW side of Rabbit Key. Syringodium and Thalassia were the dominants, and the station was representative of zone 3. Diplanthera and Halophila were also present. Water clarity was the greatest of any station. The depth varied with the tides from 1-2 m. Station 6, near the western margin of Anclote estuary, was located about 100 m off the eastern shore of Anclote Key at the southern end of Dutchman Key. Thalassia was the only seagrass and the station was typical of zone 2 but differed from station 2 in that the turbidity was greater at station 6 because of the pres- ence of fine sediments and strong tidal currents. The depth ranged, with tide, from about 0.5 to 1.3 m. The location of each station is shown in figure 1. Voucher specimens of all algal species in the annotated list which follows have been deposited in the herbarium of the Department of Marine Science, St. Petersburg Campus, University of South Florida. Discussion—Since Diplanthera is somewhat more euryhaline and eury- thermal than any other western Atlantic species of seagrass, it extends farther up salinity gradients into estuaries such as Tampa Bay and Charlotte Harbor, and also extends farther north along the Atlantic coast, at least to North Caro- lina. The other species of seagrasses in the Anclote estuary are not known to ex- tend north of Cape Canaveral on the Atlantic coast. All are basically continuous around the Gulf of Mexico except where their distribution is interrupted, at least No. 3, 1975] BALLANTINE AND HUMM—ALGAL EPIPHYTES 153 = ) TARPON TB a ; SPRINGS Fig. 1. Map of the Anclote estuary showing location of the six stations at which field studies and collections were made. The inset of the State of Florida in the upper right corner shows the area of the map by means of a square. in shallow water, by river discharge (Humm 1956). It appears that Diplanthera does not compete well with Thalassia and Syringodium in areas that are more or less optimum for the latter species. Where Thalassia and Diplanthera are mixed, the habitat is probably suboptimum for Thalassia. Where Diplanthera is in a pure stand, the environment may be subminimal for Thalassia. In the An- clote estuary along the outer margin of the seagrass beds, an outer band of Di- planthera is often found beyond the outermost Thalassia. This pattern does not occur in all seagrass areas of the Florida coast and has not often been described. Halophila engelmannii is usually found mixed with Thalassia and then rather sparsely so. However, one small area of Halophila in pure stand was found near station 4. Halophila baillonis has not been found in the Anclote estuary during the present work, although it is to be expected. Comparison with Miami. Humm (1964) recorded 113 epiphytes of Thalassia in the Miami area, with special reference to Biscayne Bay. In this report, a total of 66 species has been recorded on all seagrasses, but in a much smaller area. Of the 66 species recorded at Anclote, 51 (about 75%) were also recorded by Humm at Miami, indicating a high degree of similarity between these populations of benthic algal epiphytes on seagrasses. About 65% of the benthic algae occurring in the Anclote estuary occur as seagrass leaf epiphytes. Host Specificity. While some species of algae were found upon only one species of seagrass in this study, a careful consideration of the host-epiphyte data suggests that there is little host specificity. Myriotrichia subcorymbosa and Sticty- osiphon subsimplex appear to be much more abundant, in most areas, on Di- 154 FLORIDA SCIENTIST [Vol. 38 planthera than on any other seagrass. This observation, however, seems to be related to the fact that both these species grow best in the shallowest water oc- cupied by seagrasses where there are only pools during spring low tides rather than an adaptation to Diplanthera leaves. The deepest Diplanthera beds did not support these species. Cladosiphon occidentalis was more abundant on Thalassia than on other seagrasses, but the reason appears to be its adaptation to some- what deeper water rather than to Thalassia leaves. Most species of algae oc- curred on all species of seagrasses in proportion to the abundance of each sea- grass. ANNOTATED LIST OF EPIPHYTIC ALGAE A key to the identification of all species in the list that follows is found in Dawes’ (1974) Marine Algae of the West Coast of Florida. CYANOPHYTA Order COCCOGONALES Determinations in the order Coccogonales are based upon the revision of the coccoid Myxophyceae by Drouet and Daily (1956). Family CHROOCOCCACEAE Anacystis aeruginosa Drouet and Daily. Forming microscopic, gelatinous colonies on the older leaves of Thalassia and occasionally on Syringodium and Diplanthera at stations 1 and 5. Best developed in areas of low current velocity and low wave action, as it is readily dislodged from the leaves. Anacystis dimidiata Drouet and Daily. This species rarely forms colonies of more than a few cells and is often found as a solitary cell or one that has recently divided and the daughter cells still have flattened adjacent faces. It is microscopic, widely distributed, but found in this study on Diplanthera and Thalassia at station 1 only. Anacystis montana (Lightfoot) Drouet and Daily. Forming bright green patches on all species of seagrasses at stations 1, 2, 3, 5, and 6. This species is primarily a fresh water one but it will grow in estuaries. In the Anclote area it probably comes down the river as temporary plankton and becomes affixed to various solid surfaces, including seagrass leaves. Agmenellum thermale (Kitzing) Drouet and Daily. This is another bluegreen that occurs as plankton, attached lightly to solid surfaces, and lives in intertidal sand. In the Anclote area it was found only once on the surface of a Syringodium leaf from station 5. Since it is microscopic and requires high magnification to recognize, it is easily over- looked despite its wide distribution. Family CHAMAESIPHONACEAE Entophysalis conferta Drouet and Daily. Common at all stations and at all seasons of the year on Diplanthera, Thalassia and Syringodium attached directly to the seagrass leaves and also as epiphytes of many other algae that grow epiphytically on the sea- grasses. The colonies are microscopic. Entophysalis duesta (Meneghini) Drouet and Daily. This species penetrates lime- stone and was found at all seasons and at all stations within the calcareous tubes of serpulid worms and the tests of Bryozoa (Schizoporella) on Thalassia, Syringodium, and Diplanthera. The cells of E. deusta tend to be oval or spherical outside limestone whereas inside limestone they tend to simulate filamentous growth. In the past, the external cells have been placed in the genus Xenococcus and the internal filament-like strands in the genus Hyella (Tilden, 1910; Desikachary, 1959). No. 3, 1975] BALLANTINE AND HUMM—ALGAL EPIPHYTES 155 Order HORMOGONALES Family OscILLATORIACEAE The classification of the Oscillatoriaceae used here is that of Drouet (1968). The vari- ous synonyms or ecophene names of Drouet’s species found in the Anclote area are in- dicated by their old names in order that ready reference may be made to the older lit- erature and also in order to indicate which ecophenes of Drouet’s species were present at Anclote. Microcoleus lyngbyaceus (Kutzing) Crouan. Records for this assemblage of eco- phenes are given under the ecophene names below. Lyngbya confervoides Gomont. Producing mostly horizontal masses of long filaments with a distinct sheath on blades of Thalassia at stations 1 and 6, year around but most abundant during late summer. Lyngbya majuscula Gomont. Clusters of filaments on Thalassia and Syringodium at sta- tions 2, 3, 4, and 5. It was observed at station 5 at all seasons of the year, but only sporadi- cally at the other stations. Best development occurred in July and August. This is the largest ecophene of the species in marine waters; its trichomes reach 60 um diam or more. Lyngbya semiplena Gomont. This slender ecophene was found at all stations and at all seasons on the three common species of seagrasses, Thalassia, Syringodium, and Di- planthera. During the summer months, however, it developed in great abundance, form- ing skeins of filaments that almost covered the seagrass leaves, especially in the more shallow water stations and especially over stands of Diplanthera. It readily tolerates tem- peratures of 38° C or more. Oscillatoria lutea C. Agardh. Forming gelatinous patches on the leaves of sea- grasses at all seasons at most stations. Probably present at all stations the year around. The plant masses are usually yellow-brown or olive; the trichomes coiled or entangled in this strictly marine species, formerly Lyngbya lutea (C. Agardh) Gomont. Porphyrosiphon notarisii (Meneghini) Kitzing. This species was present as the eco- phene, Oscillatoria nigroviridis, on Thalassia at station 2 in May. It was more abundant throughout the estuary on shells and mangrove roots in shallow water. Schizothrix arenaria (Berkeley) Gomont. This species was present as two of its eco- phenes, sensu Drouet 1968, as follows: Oscillatoria laetevirens Gomont was found on Thalassia and Diplanthera at station 5 only during winter and summer, although it is probably present the year around. Its slender trichomes tend to be grouped into small cushions or pads on the grass leaves and also on bottom sediments in grass beds. Occasionally it occurs in a thin Phormidium-like mem- brane of appressed trichomes. Microcoleus chthonoplastes Thuret, while most abundant on muddy sand bottom sedi- ments in the intertidal zone also occurred on Thalassia and Diplanthera leaves at station 1 as thin, membranous layers of filaments several months during the year but especially in late winter. Schizothrix calcicola (C. Agardh) Gomont. This species, as three ecophenes, was one of the most widely distributed bluegreens in the Anclote area. It was present at all sta- tions on all species of seagrasses throughout the year. Lyngbya digueti Gomont occurred as a microscopic turf on Diplanthera and Thalassia at stations 1 and 2, probably the year around. Its filaments were attached to the hori- zontal, basal parts that were curled and entangled, but the upright parts were straight and vertical. Lyngbya epiphytica Hieronymus was most often encountered spiralling around fila- mentous epiphytes of all three common species of seagrasses, especially Polysiphonia, Ceramium, Cladophora and filamentous bluegreens. While not recorded at all stations and all seasons, it was probably present, as it is easily overlooked. Plectonema terebrans Bornet and Flahacult is an ecophene of S. calcicola that bores into 156 FLORIDA SCIENTIST [Vol. 38 limestone. It was found on Thalassia and Syringodium within the calcareous tubes or tests of serpulid worms and encrusting Bryozoa to which it usually imparts a greenish tinge. It was present at all stations the year around. Schizothrix mexicana Gomont. Two ecophenes of S. mexicana were found in the An- clote area, though neither was abundant. Lyngbya gracilis Gomont formed small tufts of filaments on all 3 seagrasses, Diplan- thera, Thalassia, and Syringodium, at all times of the year and at stations 1, 2, and 3. Though widely distributed, it was never very abundant. Lyngbya sordida, an ecophene that produces large trichomes (15-30 um diam.) was found only on Thalassia at station 1; the filaments were in fasciculate tufts. It is not typical as an epiphyte. | Spirulina subsalsa Oersted. On leaves of all seagrasses at stations 1, 2, and 5 during May and July in the form of small bright bluegreen patches. Part of the collections fitted the ecophene S. major Gomont, as the spirals were loose, the turns several microns apart. Family RivuLARIACEAE Calothrix crustacea Schousboe and Thuret. This species is often an epiphyte and occurred on all species of seagrasses at Anclote and at all stations throughout the year. It formed small, blackish tufts when well developed, but often the growth was micro- scopic. This name now includes many species in a number of genera, sensu Drouet, 1973. Some of these plants had heterocysts at the base only and would have been referred to C. confervicola (Roth) C. Agardh in the older literature. RHODOPHYTA Order BANGIALES Family BANGIACEAE Asterocystis ramosa (Thwaites) Gobi. Though not recorded for August, Asterocystis appears to be year around with a peak of abundance during spring. On all 4 species of seagrasses, and at all stations except number 3. Erythrocladia subintegra Rosenvinge. Forming microscopic red discs on Thalassia and Diplanthera during May, June, and July at station 5. It was probably present at other stations and during other months. Erythrotrichia carnea (Dillwyn) J. Agardh. The most common member of the family in the Anclote area. Present on all snecies of seagrasses at all stations the year-around. During spring, when best developed, it often forms a fine red fuzz on the margin of Di- planthera and Thalassia. Goniotrichum alsidii (Zanardini) Howe. Occasional on all 4 seagrasses and recorded once or more at stations 1, 2, 5, and 6 every month except January and May. Probably present the year around. Order NEMALIONALES Family ACROCHAETIACEAE Acrochaetium sargassi Borgesen. On all 4 seagrasses at all stations during January, March, April, July, and August. Although this species was originally described by Bérge- sen on Sargassum in the Virgin Islands, it is known from a variety of algae. It was reported for Tampa Bay on Thalassia by Dawes (1967). Acrochaetium thureti (Bornet) Collins and Hervey. On Thalassia and Diplanthera at stations 1 and 2, in May. This species has been reported by Taylor (1957, 1960) only from Massachusetts and Bermuda. Aziz (1965), however, found it at Beaufort, N. C., on Gelidium crinale and in Biscayne Bay, Miami, on Caulerpa prolifera. Acrochaetium crassipes Borgesen. On Diplanthera at stations 2 and 3, April and May. This warm water species was classified as Kylinia crassipes (Borgesen) Kylin by Taylor (1960, p. 300) but we follow the opinion of Aziz (1965) who interpreted the genus Acro- chaetium in the sense of Rosenvinge (1909). No. 3, 1975] BALLANTINE AND HUMM—ALGAL EPIPHYTES 157 Order CRYPTONEMIALES Family SQUAMARIACEAE Peyssonnelia rubra (Greville) J. Agardh. On Syringodium at station 2, March. This species is usually on stones or shells and is usually well below low tide. Family CORALLINACEAE Fosliella atlantica (Foslie) Taylor. Abundant at all seasons and all stations on all spe- cies of seagrasses, at times almost completely covering the leaves, especially of Thalassia. This calcareous, encrusting epiphyte promotes the attachment of other epiphytic algae by offering a better substratum than the seagrass leaves themselves. Fosliella farinosa (Lamouroux) Howe. Perhaps as abundant as F. lejclisii and as widely distributed, but here recorded on Thalassia and Diplanthera at stations 1 and 2 during July and August. This plant is often 2 or 3 cells thick and has colorless, swollen cells (trichocytes) terminating the cell rows. These 2 species probably reduce significantly the photosynthesis on the seagrass leaves when present in abundance. Jania adhaerens Lamouroux. Mostly on the basal portion of leafy branches of all 4 seagrasses, but also on their leaves, at stations 1, 2, 3, and 4. Recorded for January, Febru- ary, March, May, and July, but probably present the year around. Family HyPNEACEAE Hypnea musciformis (Wulfen) Lamouroux. Occasional in the form of small plants on all 3 kinds of seagrasses the year around at stations 2, 3, 4, 5, and 6, and sometimes station 1. These plants apparently break off the seagrass leaves before they get very large in re- sponse to periods of windy weather. Drifting plants often reattach to seagrass leaves by means of the hooked tips on the branches that have a tendency to enwrap anything they come in contact with very promptly if it is small enough for them to surround. Thus the abundance of Hypnea on seagrass beds varies with weather conditions. Large quantities may accumulate during periods of calm weather, most of which is washed ashore or out to sea during periods of windy weather. Hypnea grows rapidly during warm weather. It produces an abundance of kappa carrageenan as the principal cell wall constituent and for this reason is of considerable economic value. Hypnea spinella (C. Agardh) Kitzing. This small, slender species is uncommon at Anclote, having been found on Thalassia, Syringodium, and Diplanthera at stations 2, 5, and 6 only during July and August. Order RHODYMENIALES Family CHAMPIACEAE Champia parvula (C. Agardh) Harvey. Common on all 4 seagrasses at stations 1, 2, 3, 4, and 6 from January to April and in July and August. Order CERAMIALES Family CERAMIACEAE Centroceras clavulatum (C. Agardh) Montagne. A common epiphyte on all 4 species of seagrasses the year around at all stations. Ceramium byssoideum Harvey. Typical plants of this species were not common but occurred on Diplanthera at stations 1, 2, and 4. What is considered a form of this species was very common. It is described below. 158 FLORIDA SCIENTIST [Vol. 38 Fig. 2. Photograph of Ceramium byssoideum forma alternata showing the alternate branching that distinguishes this form from the species in which the branching is dichotomous. Ceramium byssoideum forma alternatum forma nova C. byssoideum byssoideum persimilis, sed ramulis alternatis differt. This form was similar to the species in all major respects including nodal cortication and various dimensions but differed in having distinctly alternate branching rather than di- chotomous branching with the associated forcipate tips. This newly described form was found the year around on all 4 species of seagrasses. The type specimen has been de- posited in the herbarium of the Department of Marine Science, University of South Florida, St. Petersburg (Accession number 974). Ceramium fastigiatum (Roth) Harvey, forma flaccidum Peterson. This species was common and recorded from all 4 seagrasses at all months during which collections were made except January and March at stations 1, 2, and 5. Griffithsia tenuis C. Agardh. Found only 3 times, during May, June, and July at sta- tions 2 and 5 on Syringodium only. Griffithsia globulifera Harvey. Found only once at station 2 on Thalassia in Febru- ary. Spyridia filamentosa (Wulfen) Harvey. One of the most common red algal epiphytes throughout the year at all stations and on all species of seagrasses. This species often orig- inates on seagrass leaves but is torn loose by windy weather before the plants are very large. Like Hypnea, Spyridia also develops hooked tips at the apex of some branches and these have the ability to enwrap seagrass leaves, especially Syringodium and Diplanthera, or other algae when they come in contact for a few hours. Thus drifting plants may be- come reattached to seagrasses. This species also readily produces adventitious attach- ment organs from any axis in response to contact of sufficient duration. The peak of abundance is reached during late spring. Family DasyACcEAE Dasya pedicellata (C. Agardh) C. Agardh. Found only once in the April collection on Syringodium from station 2, a small, sterile plant. Family RHODOMELACEAE Polysiphonia havanensis Montagne. Occasional on all seagrasses at stations 1, 2, and © during April, May, July, and August. Best developed in spring. Polysiphonia echinata Harvey. One of the most common red algal epiphytes, recorded from all species of seagrasses at all times of the year but best developed during March and April. During the warmer months of the year the main axes of the plants were mostly 70-100 um in diam., but in winter and spring they were about 150 um. Herposiphonia tenella (C. Agardh) Ambronn. Recorded during July and August on Diplanthera, Thalassia, and Syringodium at stations 1, 5, and 6. This plant has a creep- ing main axis with erect, determinate branchlets. No. 3, 1975] BALLANTINE AND HUMM—ALGAL EPIPHYTES 159 Lophosiphonia saccorhiza Collins and Hervey. Recorded on all 4 seagrasses January through August at stations 1, 2, 5, and 6. This species also has a creeping main axis and erect, determinate branchlets, the tips of which are curved, with trichoblasts on the con- vex side. Humm (1964) comments on the uncertainty of this determination. Chondria collinsiana Howe. This small species is typically an epiphyte. It was found from January through August at stations 1, 2, 4, 5, and 6 on all 4 species of seagrasses. It was not recorded for Tampa Bay by Dawes (1967). Chondria dasyphylla (Woodward) C. Agardh. This and the following species are rarely epiphytes but occasionally attach to seagrass leaves. Small plants were found a few times at stations | and 5 in January, May, and July on all 4 seagrasses. Larger plants break loose or, in the case of old seagrass leaves, cause the entire leaf to break off. Chondria tenuissima (Goodenough and Woodward) C. Agardh. Found only once on Diplanthera at station 2 in the January collection. Laurencia obtusa (Hudson) Lamouroux. Found once on Diplanthera during January at station 2. Laurencia poitei (Lamouroux) Howe. Found occasionally on Thalassia and Syringo- dium, at stations 2, 5, and 6 in February and during spring and summer. This and the previous species are rarely epiphytes and do not grow to maturity on seagrass leaves. PHAEOPHYTA Order ECTOCARPALES Family ECTOCARPACEAE Giffordia rallsiae (Vickers) Taylor. Occasional during February and April at station 2 on Diplanthera and Syringodium. This is a tropical member of the genus known throughout the West Indies and Caribbean Sea, so its presence only during winter and spring is puzzling. Giffordia mitchellae (Harvey) Hamel. Abundant during winter and spring, but pres- ent the year around (except during July?) at all stations and on all species of seagrasses. This is a temperate species with a wide latitudinal distribution from the tropical Atlantic to Nova Scotia. In its southern range it is most abundant during winter and spring, but in its northernmost range it is a plant of summer and fall. Order SPHACELARIALES Family SPHACELARIACEAE Sphacelaria furcigera Kitzing. Collected during spring and summer only on Tha- lassia and Syringodium at stations 2, 5, and 6. It occurred around the base of the upright branches of the seagrass and not on the leaves. Sphacelaria tribuloides Meneghini. On basal parts of the upright branches of Thalassia and Syringodium from stations 2 and 6 during March, May, and July. It was probably present the year around. Order CHORDARIALES Family CHORDARIACEAE Cladosiphon occidentalis Kylin. This species appeared on Thalassia in December, was most abundant on this seagrass and occasionally on the other 3 at most stations until early May when it disappeared. During winter and spring it was the most conspicuous brown alga in the area. The form in which it is present during the warmer months is not yet known. Family MyYRIONEMATACEAE Ascocyclus magnusii Sauvageau. Common on Thalassia at station 2 during March. It was probably widely distributed during winter and spring, but apparently not present during ‘the summer months. This species forms microscopic discs on the leaf surface. Known from Florida to New England but in its northern range it is a plant of spring and summer. Taylor (1960) and Earle (1969) list it as A. orbicularis Magnus. 160 FLORIDA SCIENTIST [Vol. 38 Order DicTYOsIPHONALES Family SticryOsIPHONACEAE Myriotrichia subcorymbosa (Holden) Blomquist. Abundant on Diplanthera, especially in the shallow water stations 1 and 2 the year around; occasional on Thalassia and Syrin- godium. It is often the most common epiphyte on Diplanthera, forming a fine brown fuzz or fringe. Although it is abundant throughout the West Indies and around the entire Florida coastline, Taylor (1960) reports it for North Carolina only and (1957) from Mary- land to Massachusetts, both as Ectocarpus subcorymbosus Farlow emend. Holden. Stictyosiphon subsimplex Holden. Abundant on all 3 seagrasses, but especially on Di- planthera, during winter (from November) and spring (until early May) at all stations but especially at the shallow water stations just below mean low tide. This species occurs from the West Indies to Nova Scotia. In its southern range it is present during winter and spring; in its northern range it is ephemeral, and present during the fall or during spring and fall, almost exclusively on the fresh and brackish water grass, Ruppia. Taylor (1957) reported it for Connecticut and southern Massachusetts only; it is not listed in his 1960 publication. Fiore (1970) has shown that this species is the sporophyte and Myrio- trichia subcorymbosa the gametophyte of a plant that exhibits a life history influenced by environmental conditions. He has proposed a name change for this plant in a forth- coming publication. Earle (1969, figs. 98, 105, 107) provides excellent illustrations of this species and discusses its distribution in the Gulf of Mexico. CHLOROPHYTA Order ULOTRICHALES Family PLEUROCOCCACEAE Pseudotetraspora antillarum Howe. Found from January to May at stations 1, 2, and 5 on all 4 species of seagrasses. Young plants frequently appeared to become estab- lished on the seagrass leaves at the vortex of the spirals of calcareous tubes (about 1 mm diam.) of a serpulid worm. Family GOMONTIACEAE Gomontia polyrhiza (Lagerheim) Bornet and Flahault. Creeping on the surface of the leaves of Thalassia and Diplanthera and penetrating empty surface cells of the host. The filament of the alga is markedly constricted at the point of penetration of the cell and the cells are generally irregular, reaching a maximum diam. of 20 wm, considerably greater than filaments that penetrate limestone, a common characteristic of this species. It was found during July only at stations 5 and 6, but since it is a microscopic species easily over- looked, it may be present at Anclote the year around. While previously reported as pene- trating wood (Bornet and Flahault, 1888), apparently this is the first report of this species penetrating seagrass leaves. Family CHAETOPHORACEAE Entocladia viridis Reinke. This is another microscopic species that creeps within the surface polysaccharide of a great variety of larger algae. Though recorded only during July as an epiphyte of Polysiphonia echinata from a Thalassia leaf, it is very likely that it is a year-around alga in the surface layer of larger species at all stations and from all species of seagrasses. Phaeophila dendroides (Crouan) Batters. Found upon the leaves of all 4 seagrasses and occasionally upon larger algae that were seagrass epiphytes; stations 1, 2, 5, and 6 in the July and August collections, but probably the year around. This microscopic species, though very common, is easily overlooked. No. 3, 1975] BALLANTINE AND HUMM—ALGAL EPIPHYTES 161 Ulvella lens Crouan. Recorded on Thalassia leaves at stations 1 and 2 during May only. This species is also microscopic, easily overlooked, and probably occurs on all species of seagrasses at all stations the year around. Order ULVALES Family ULVACEAE Enteromorpha chaetomorphoides Borgesen. Although this is a tropical species, it was recorded during January and February only at station 4 on Thalassia and Diplanthera but wrapped around or entangled among the seagrasses rather than attached. Enteromorpha clathrata (Roth) J. Agardh. Although this is a common, year-around species in the area, it was found epiphytic only on Thalassia and only during April at sta- tion 4. Enteromorpha prolifera (Miller) J. Agardh. Another year around and abundant spe- cies that reaches a peak of development during winter and spring. It was recorded as an epiphyte, however, only during January at stations 1 and 4 on Diplanthera, Thalassia, and Syringodium. A number of juvenile Enteromorpha plants were not recorded because of the lack of critical characters that would permit reasonably confident determinations. Order CLADOPHORALES Family CLADOPHORACEAE Chaetomorpha brachygona Kitzing. Though not found attached to seagrasses it was present entangled among the leaves at all seasons at stations 2 and 5. Cladophora sericea (Hudson) Kitzing. Small plants were common the year around but most abundant during winter and spring at all stations except number 3 and on all species of seagrasses. These plants would mostly fall within the range of C. gracilis, C. glaucescens, or C. flexuosa sensu Collins (1909) and Taylor (1957, 1960). The mono- graph of van den Hoek (1963) has been followed in this determination. Rhizoclonium hookeri Kitzing. Found on Syringodium and Halophila at station 2 in January and February. Rhizoclonium kerneri Stockmayer. Common from May to August and probably during the fall on all species of seagrasses at stations 1, 2, 3, 5, and 6. This almost microscopic plant often forms a green fuzz on seagrass leaves. Order SIPHONALES Family DERBESIACEAE Derbesia vaucheriaeformis (Harvey) J. Agardh. Found during all seasons on Halophila, Syringodium, and Diplanthera at station 5 only. Halicystis osterhoutii, said to be another form of this plant, has not been reported along the Florida Gulf Coast. It may occur on rocky areas off shore in deep water. Caulerpa prolifera (Forsskal) Lamouroux. While not an epiphyte of a seagrass in the same sense as other species listed in this paper, a small plant of C. prolifera was collected attached to the basal portion of an erect branch of Thalassia at station 5 in May. A cluster of rhizoids at the base of the plant held a clump of sediment but the plant was not at- tached to the bottom. The Caulerpa zygote or “seed” material may have originally lodged upon the Thalassia plant. ACKNOWLEDGMENTS—Grateful acknowledgment is made to Florida Power Corporation of St. Petersburg for a graduate research stipend and expense funds that made this work possible. 162 FLORIDA SCIENTIST [Vol. 38 LITERATURE CITED Aziz, K. M. S. 1965. Acrochaetium and Kylinia in the Southwestern North Atlantic Ocean. Ph.D. Dissert. Duke Univ. Durham, N.C. Bairp, R. C. (ed.) 1972. Anclote Environmental Project Report for 1971. Dept. Marine Sci. Univ. S. Florida. pp. 251. (Mimeographed) BorneT, E., anp C. FLaHavuLt. 1888. Note sur deux nouveaux genres d’algues perforantes. J. de Bot. 2:161-165. Co..ins, F. S. 1909. The green algae of North America. Tufts College Stud. (Sci. Ser.) 2(3):79-480. Dawes, C. J. 1967. Marine algae in the vicinity of Tampa Bay, Florida. Privately reproduced. pp. 150+9 pl. _______. 1974. Marine Algae of the West Coast of Florida. Univ. Miami Press. Coral Gables. Desikacuarky, T. V. 1959. Cyanophyta. Indian Council of Agricultural Research. New Delhi. DrovEt, F., anp W. A. Dairy. 1956. Revision of the coccoid Myxophyceae. Butler Univ. Bot. Stud. 12:1-218. . 1968. Revision of the classification of the Oscillatoriaceae. Acad. Nat. Sci. Philadelphia Monogr. 15:1-370. . 1973. Revision of the Nostocaceae with Cylindrical Trichomes. Hafner Publ. Co. New York. Earte, Syivia A. 1969. Phaeophyta of the eastern Gulf of Mexico. Phycologia 7:71-254. . 1972. Benthic algae and seagrasses. In BUSHNELL, V. C. (ed.). Serial Atlas of the Marine Environment. Folio 22:15-18 + pl. 6. Amer. Geogr. Soc. New York. FEIcL, J. L., anp T. E. Py.e. 1973. Application of aerial photography to the study of seagrass beds and turbidity in a Florida estuary. In Anclote Environmental Report for 1972. Dept. Marine Sci. Univ. S. Florida. Appendix 3B: 199-204. (Mimeographed) Fiore, J. 1970. Life History Studies of Phaeophyta from the Atlantic Coast of the United States. Ph.D. Dissert. Duke Univ. Durham, N.C. HokExk, C. vAN DEN. 1963. Revision of the European Species of Cladophora. E. J. Brill. Leiden. Huma, H. J. 1956. Sea grasses of the northern Gulf coast. Bull. Mar. Sci. 6:305-308. . 1964. Epiphytes of the sea grass, Thalassia testudinum, in Florida. Bull. Mar. Sci. 14:306- 341. . 1973. Seagrasses. In Jones, J. I. et al. (eds.). A Summary of Knowledge of the Eastern Gulf of Mexico. pp. IIIC-1 to IIIC-10. Amer. Petrol. Inst. Washington, D.C. , AND SYLVIA EARLE Tay_or. 1961. Marine Chlorophyta of the upper west coast of Florida. Bull. Mar. Sci. 11:321-380. Jouansson, R. J. O., ano T. L. Hopkins. 1973. Primary productivity. In Anclote Environmental Report for 1972. Dept. Marine Sci. Univ. S. Florida pp. 48-59. (Mimeographed) Pyxe, T. E., J. C. McCartuy, anp G. M. Grirrin. 1973. Reconnaissance mapping of turbidity in estuaries. In Anclote Environmental Report for 1972. Dept. Marine Sci. Univ. S. Florida. Appendix IB: 189-196. (Mimeographed) RoseEnvINGE, L. K. 1909. The marine algae of Denmark. Part I, Rhodophyceae I. Kgl. Dansk Vidensk. Selsk. Skrifter, Nat. og. Mathem, Af. 7, Raekke, 7(1):1-151. TayLor, W. R. 1957. Marine Algae of the Northeastern Coast of North America. Rev. ed. Univ. Michigan Press. Ann Arbor. . 1960. Marine Algae of the Eastern Tropical and Subtropical Coasts of the Americas. Univ. Michigan Press. Ann Arbor. TILDEN, J. E. 1910. Minnesota Algae, Vol. I. The Myxophyceae of North America and Adjacent Re- gions Including Central America, Greenland, Bermuda, the West Indies, and Hawaii. Univ. Minn. Bot. Ser. 8: 1-328. ZIMMERMAN, R. J., R. A. Dietz, T. E. Pte, S. W. Rocers, N. J. BLAKE, AND H. J. Hum. 1973. Ben- thic community—seagrasses. In Anclote Environmental Report for 1972. Dept. Marine Sci. Univ. S. Florida. pp. 115-141. (Mimeographed) Florida Sci. 38(3):150-162. 1975. Biological Sciences ELEMENTAL ANALYSIS OF SELECTED MERRITT ISLAND PLANTS Davip H. Vickers, ROSEANN S. WHITE, AND I. JAck STOUT Department of Biological Sciences, Florida Technological University, Orlando, Florida 32816 ABSTRACT: Samples of mature leaves from various plant species were collected from experi- mental plots located at Kennedy Space Center during August 1973 and April 1974. Concentrations of the following elements were measured in each of the samples using atomic absorption techniques: K, Na, Ca, Mg, Fe, Mn, Zn, Cu, and Al. The concentrations of B and Mo were found to be below the sensitivity of the method. ELEMENTAL composition of selected plant species at Kennedy Space Center was determined to establish base levels so that future studies could be initiated to discover whether solid fuel rocket launchings cause perturbations in the quan- tities of elements commonly recognized as essential for normal plant growth and development. The complexity of comparative mineral nutrition of plants has been reviewed extensively by Gerloff (1963). The 10 primary elements essential to all plants are: carbon (C), hydrogen (H), oxygen (O), nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), sulfur (S), magnesium (Mg) and iron (Fe). At least 5 other elements, manganese (Mn), zinc (Zn), boron (B), copper (Cu), and molybdenum (Mo) are required in minute amounts for the growth of most plants and sodium (Na), aluminum (Al), silicon (Si), chlorine (Cl), gallium (Ga), va- nadium (V), and cobalt (Co) have been shown to be essential for the normal growth of some species of plants. However, not all species require the same ele- ments or the same concentrations of elements, and some species even accumulate elements to concentrations that are toxic to other plant species. MATERIALS AND METHops—Plant material was collected from two areas on north Merritt Island, Florida. Sample areas 1-8 are located on a poorly drained site with a hardpan and seasonal flooding is typical. The plant community is similar to that found over much of Florida and referred to as pine flatwoods (Edmisten, 1963) but the community sampled is unique in not having a pine over- story. Sample areas 9-16 are located on a deep, well-drained soil which does not flood. The plant community is best characterized as scrubby flatwoods (Laessle, 1942:29). Smith (1962) reviewed mineral analysis of plant tissue and determined that: a) leaf samples most accurately reflect the elemental composition of the whole plant; and b) the elemental content of mature leaves is less in a state of flux than that of young leaves. He reported that 25 selected leaves from a citrus tree will give a valid measure of the elemental composition of the entire plant. We accepted his findings and sampled accordingly. Mature terminal leaves from each of the chosen plant species were collected in August 1973 near the end of the rainy season. Foliar leaching should have 164 FLORIDA SCIENTIST [Vol. 38 equilibrated by this time. Species sampled included saw palmetto (Serenoa repens), Quercus minima, Q. pumila, Q. myrtifolia, and Q. chapmanii. Limited samples were taken of wax myrtle (Myrica cerifera), Ilex glabra, and Lyonia fru- ticosa, since they did not occur on all experimental plots. Three of the species (S. repens, Q. chapmanii, and Q. myrtifolia) were also sampled in April of 1974, as was the additional species, Q. virginiana. Leaf material in the April sample had been set at the end of the dry season and probably little foliar leaching had occurred. Leaves were picked with new disposable plastic gloves and frozen until analyzed. The samples were dried at 110°C for 24 hr and weighed to the nearest mg. The method of Kometani, et al. (1972) was used to ash plant samples for atomic absorption analysis. Tissues were placed in acid washed borosilicate beakers and heated at 300°C for 30 min in order to prevent flaming and loss of matter which results from heating at higher temperatures. The samples were then heated at 550°C for 1 hr. This procedure reduced the plant tissues to an inorganic ash which was extracted twice with w/w nitric acid. The extract was heated almost to dryness and brought up to a final volume of 10 ml with distilled water. The concentrations of 11 elements (K, Na, Ca, Mg, Fe, Mn, Zn, Cu, Al, B, Mo) were measured in each of these samples using a Perkin-Elmer Model 303 Atomic Absorption Unit (Perkin-Elmer Corp., 1966). REsuLTs—A summary of all data is presented in Table 1. The two columns in- dicate the August 1973 and April 1974 sampling dates respectively. The most extensive data exist for saw palmetto. In all cases the concentrations of boron and molybdenum were below the lower limits of detection for the method employed, i.e., 22 and 2 ppm respectively. Data presented in Table 1 follow the same general pattern reported for other plant tissues. The majority of the cations present in all of the 9 species analyzed could be accounted for by Na, K, Ca, and Mg. The extreme variation within species makes it almost impossible to make any definite statements as to seasonal variations and differences among plots of the same species. The atomic absorp- tion unit used for these determinations is +5% accurate and if one allows an- other +5% for weighing and dilution errors, a total variance of + 10% might occur. In many cases, the variance greatly exceeded 10%. One can conclude that considerable variation occurs naturally from plot to plot and from plant to plant. Seasonal variations are also to be considered (Woodwell, 1974). For example, data for Serenoa repens clearly show that the potassium concentration of spring growth (April 1974) is significantly greater than late summer growth (August 1973); while sodium exhibits the opposite behavior. Two species (Serenoa repens and Quercus minima) were collected exten- sively over 16 experimental plots representative of pine flatwoods (plots 1-8) and scrubby flatwoods (plots 9-16). Data from the two areas were compared with no significant differences found. It would appear the physical location of these species did not significantly affect mineral content. In order to see species differences element by element more clearly, data presented in Table 1 have been plotted on Fig. 1-9 as ppm of a given element No. 3, 1975] VICKERS ET AL.—ELEMENTAL ANALYSIS OF PLANTS 165 Tas_e 1. Elemental concentration (ppm/g dry tissue) of leaf tissue from selected plant species collected on North Merritt Is- land, Florida. POTASSIUM SODIUM CaLcium MAGNESIUM SPECIES Aug.* Apr. Aug. Apr. Aug. Apr. Aug. Apr. A. Serenoa repens 5100 9100 3800 2400 830 1100 1500 1440 std. dev. + 1200 +1900 + 1700 +840 +230 +270 +410 +310 B. Ilex glabra 4300 - 2400 - 2700 - 970 - C. Lyonia fruticosa 5600 - 870 - 6300 - 1500 - std. dev. +3500 +330 +2700 +270 D. Myrica cerifera 2800 - 3400 - 10,000 = 7000 = std. dev. +610 +310 + 1600 +1200 E. Quercus chapmanii 8100 8700 2600 420 9900 5300 3800 1600 std. dev. +2300 + 1800 + 1600 +82 +4600 +950 +2100 +1530 F. Q. minima 5700 - 2600 - 8000 - 2300 - std. dev. +2400 +4300 +3400 + 1300 G. Q. myrtifolia 5100 6200 900 530 7200 4300 2600 1600 std. dev. +1100 +1100 +400 +180 +3500 +790 +560 +74 H. Q. pumila 5200 - 800 - 18,000 - 2400 - std. dev. + 2500 +270 +7000 +70 I. Q. virginiana - §300 760 - 2200 - 1700 std. dev. +450 +230 +390 +100 IRON MANGANESE ZINC CoprerR ALUMINUM Aug.* Apr. Aug. Apr. Aug. Apr. Aug. Apr. Aug. Apr. A. Serenoa repens 19 23 16 30 7.7 13 1.9 3.7 5.0 7.8 std. dev. +6.8 +6.9 +3.1 +9.6 +2.8 +3.3 + .82 +1.2 +91 +3.2 B. Ilex glabra 3.9 - 43 - 24 - 8.8 - 100 - C. Lyonia fruticosa 6.4 - 43 - 21 - 7.0 - 61 - std. dev. +1.8 +27 +9 +3.2 +34 D. Myrica cerifera 6.6 - 34 - 27 - 6.2 - 87 - std. dev. +14 +11 +5.5 +1.4 +26 E. Quercus chapmanii 74 55 ttl 64 22 19 5.2 6.2 47 24 std. dev. +15 +11 +26 +31 +2.7 +2.8 +1.3 +1.4 +18 +62 F. Q. minima 69 - 55 - 26 _- 8.7 - 120 - std. dev. +40 +25 +7.3 +9.6 +190 G. Q. myrtifolia 64 53 100 140 29 30 4.1 5.4 51 35 std. dev. +26 +21 +16 +26 +3.0 +6.0 +1.0 + .53 +8.0 +14 H. Q. pumila 8.5 - 41 - 31 - 6.9 - 81 - std. dev. +6.4 +33 +.71 0 +2.8 I. Q. virginiana - 63 - 104 - 34 - 11 - 41 std. dev. +8.7 +140 +5.4 +2.8 +15 *Samples from August, 1973 and April, 1974. per g dry tissue versus plant species. The figures are self explanatory so only points of special interest are discussed below. Potassium (Fig. 1). As already noted, potassium content of palmetto tissues is much higher in April than in August. All 5 oak species included in this study contained 5,000 to 9,000 ppm potassium. As in palmetto, the concentration of K was higher during April than in August for Q. chapmanii and Q. myrtifolia. My- rica cerifera contained the least amount of potassium of all 9 species analyzed. Sodium (Fig. 2). Sodium was less abundant during April than in August in S. repens, Q. chapmanii, and Q. myrtifolia in opposition to K increases. Myrica cerifera, which contained the least amount of K, contained one of the highest Na contents. The oaks, as a group, were about the same as Lyonia fruticosa with respect to Na content, being only about 1/3 to 1/4 that of palmetto. Quercus chapmanii contained much more Na during August 1973 than did the rest of the oaks but by April 1974 had fallen to a very low value. 166 FLORIDA SCIENTIST [Vol. 38 11000 9000 7000 TISSUE 5000 3000 | 1000 PPM PER GRAM SIRECIES Fig. 1. Potassium content of leaves of native plants of Kennedy Space Center. Species code is provided in Table 1. Samples A-I were collected in August, 1973 and samples A’-I’ in April, 1974. Data are shown as means + one standard deviation. Calcium (Fig. 3). Palmetto contained very small amounts of calcium at all times, whereas the other plant species as a group had significantly higher levels. Quercus pumila had the highest Ca content of all 9 species. One oak species (O. chapmanii) showed a decrease in Ca during the spring, while a second (O. myrtifolia) showed an increase. The observed seasonal changes, however, were not as great as those for Na and K. Magnesium (Fig. 4). Magnesium plays a critical role in plant metabolism as it is required for chlorophyll synthesis as well as in many essential enzymatic systems. The Mg content of palmetto was about the same during August and 5000 4000 3000 2000 1000 PPM PER GRAM TISSUE E &’ F F’ G c’ H H’ ' ’ A a’ B 8’ c c’ o D SREGIES Fig. 2. Sodium content of leaves of native plants of Kennedy Space Center. See Fig. 1 for ex- planation. No. 3, 1975] VICKERS ET AL.—ELEMENTAL ANALYSIS OF PLANTS 167 25,000 20,000 15,000 10,000 5,000 PPM PER GRAM TISSUE SPECIES Fig. 3. Calcium content of leaves of native plants of Kennedy Space Center. See Fig. 1 for ex- planation. April. Quercus chapmanii and myrtifolia both showed significant increases in Mg during August as compared to April although the variability in tissue con- centrations was much less during April than in August. Myrica cerifera averaged 7,000 + 1,200 ppm Mg, which was much higher than the other 8 plant species. Most other species ranged about 2,000 ppm Mg. Iron (Fig. 5). The oaks, with the exception of Q. pumila, were considerably higher in Fe content than the 4 other species tested. Seasonal variations from 7000 6000 5000 4000 3000 ISS 2000 1000 PPM PER GRAM SPECIES Fig. 4. Magnesium content of leaves of native plants of Kennedy Space Center. See Fig. 1 for ex- planation. 168 FLORIDA SCIENTIST [Vol. 38 August to April seemed to be minor, although decreases were observed during April as compared to August for 2 oak species. Ilex glabra, Lyonia fruticosa, and Myrica cerifera contained very little iron, possibly indicating poor uptake of this element by these plants or an ability on their part to survive on limited amounts. lu mu a60 n = 50 = G 6 n La 5 = x 4 faved © 3 faved ee) = de = a a A a’ 8 B’ c Ge D D’ E E’ F F’ G co’ H H ' v SPECIES Fig. 8. Copper content of leaves of native plants at Kennedy Space Center. See Fig. 1 for explana- tion. 170 FLORIDA SCIENTIST [Vol. 38 in August as compared to April and one increasing. The changes, in either case, were not statistically significant. Copper (Fig. 8). As was the case with Zn, palmetto contained the least amount of Cu. Most of the other species contained 5-7 ppm Cu. Quercus virginiana con- tained 11 ppm Cu—the highest for any species. Copper content was higher in April than in August for all 3 species studied over both these time periods. Aluminum (Fig. 9). Aluminum is of particular interest to us because it is the major combustion product of solid rocket fuel. Serenoa repens contains very little Al in comparison with the rest of the elements and appears to contain about the same amount in April as in August. Quercus chapmanii and myrtifolia, on the other hand, contain more Al later in the growing season (August) than during early summer (April). Myrica cerifera, Ilex glabra and Quercus pumila all con- tained more than 80 ppm Al. It appears that the single monocot (Serenoa re- pens) is able to exclude or otherwise reduce its Al uptake more effectively than the remaining 8 dicot species. 120 ‘ | PPM PER GRAM TISSUE SPEGIES Fig. 9. Aluminum content of leaves of native plants at Kennedy Space Center. See Fig. 1 for ex- planation. Discusston—Data obtained in the present study provide a baseline upon which to evaluate future studies of the plants of the area under investigation. Ex- treme variability was indicated in the mineral content of the plants encountered in the present study. Certain correlations were, however, noted: Palmetto is quite low in Ca, Zn, Cu, and Al with respect to the other plants. Based on our data, K concentrations appear to be highest in the spring and lowest during the fall; while Na follows an inverse relationship. Myrica cerifera had the lowest K content of all 9 species studied and one of the highest Na and Ca con- tents. Myrica cerifera also contained the greatest amount of Mg of all species studied. Although the oaks as a group contained relatively high concentrations of Fe and Mn, Q. pumila contained the least amount of these two elements No. 3, 1975] VICKERS ET AL.—ELEMENTAL ANALYSIS OF PLANTS 171 among the 5 oak species studied. At the same time, Q. pumila contained the highest concentration of Ca of all 9 species of plants and one of the higher Al contents. Our observations suggest that the concentration of one element should not be studied in isolation, and that consideration must be given to how the presence or absence of one element relates to other elements. ACKNOWLEDGMEN7S—The authors thank Ms. Carole Hall for technical sup- port in preparation of some of the samples for atomic absorption. The study was made possible only through the financial support and facilities provided by the National Aeronautics and Space Administration, Kennedy Space Center (Grant No. NGR 10-019-009) to Florida Technological University. LITERATURE CITED EpmisTEN, J. A. 1963. The Ecology of the Florida Pine Flatwoods. Ph.D. Dissertation. Univ. Flor- ida. Gainesville. Ger.orr, G. C. 1963. Comparative mineral nutrition of plants. Ann. Rev. Pl. Physiol. 14:107-124. Kometanl, T. Y., J. L. Bove, B. NatHANson, S. SIEBENBERG, AND M. Macyar. 1972. Dry ashing of airborne particulate matter on paper and glass fiber fitters for trace metal analysis by atomic absorption spectrometry. Environ. Sci. Technol. 6:617-620. LagssLe, A. M. 1942. The plant communities of the Welaka area. Univ. Florida Pub. 4(1):1-143. PERKIN-ELMER Corp. 1966. Analytical Methods for Atomic Absorption Spectrophotometry. Perkin- Elmer Corp. Norwalk, Conn. Situ, P. F. 1962. Mineral analysis of plant tissue. Ann. Rev. Pl. Physiol. 13:81-108. WoopweLL, G. M. 1974. Variation in the nutrient content of leaves of Quercus alba, Quercus coccinea, and Pinus rigida in the Brookhaven Forest from bud-break to abscission. Amer. J. Bot. 61:749-753. Florida Sci. 38(3):163-171. 1975. Biological Sciences RANGE EXTENSIONS FOR, AND AN ABNORMALITY IN, SCORPAENID FISHES COLLECTED OFF THE CAROLINAS WixuiaM D. ANDERSON, JR., JAMES F. MCKINNEY, AND WILLIAM A. ROUMILLAT Grice Marine Biological Laboratory, College of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412; and Department of Vertebrate Zoology (Fishes), National Museum of Natural History, Washington, D. C. 20560 ABSTRACT: Specimens of Pontinus nematophthalmus, Scorpaenodes tredecimspinosus, and Ectre- posebastes imus taken in the western North Atlantic off the Carolinas represent northeastward ex- tensions of the known ranges of these species. A specimen of Scorpaena agassizi captured off Cape Lookout, North Carolina, shows abnormalities of the dorsal fin and supporting structures. RECENT collections of scorpaenid fishes off North and South Carolina have provided specimens which extend the known ranges of Pontinus nematophthal- mus (Gunther)—the spinythroat scorpionfish, Scorpaenodes tredecimspinosus (Metzelaar)—the deepreef scorpionfish, and Ectreposebastes imus Garman and a specimen of Scorpaena agassizi Goode and Bean—the longfin scorpiontish—with abnormalities of the dorsal fin and supporting structures. 172 FLORIDA SCIENTIST [Vol. 38 A specimen (62 mm SL) of Pontinus nematophthalmus was caught southeast by south of Cape Lookout, North Carolina (34°07.7’N, 76°10.5’W, depth 71 to 143 m, R/V Eastwarp, field no. GMBL 73-65, 2050-2220 hrs, 15 May 1973, by trawl). Eschmeyer (1969) reported the distribution of this species as Florida (R/V OREGON sta. 5301), the Bahamas, throughout the Caribbean, and south to about the Amazon off Brazil in depths of about 82 to 411 m. The collection of the specimen off Cape Lookout represents an extension of the known range of the species of approximately 840 km northeastward along the Atlantic coast of the United States from R/V OREGON station 5301 at 27°18’N, 79°57’W. Because we have seen no published descriptions of live coloration for P. ne- matophthalmus or of preserved material with appreciable remnants of live color- ation, we are including notes made on our specimen 10 days after preservation. Four “poorly defined saddles” of dusky pigment present beneath dorsal fin (as noted by Eschmeyer, 1969). Body with two orange bars—first beginning near base of pectoral fin and slanting obliquely backward and disappearing at saddle below posterior part of spiny dorsal fin, second beginning above anterior part of anal fin and running almost vertically to disappear in saddle of dusky pigment below soft dorsal fin. Caudal fin with about four orange bars on proximal two-thirds of fin—the posterior three anastomosing. Considerable orange pigment at base of spiny dorsal fin—continuing onto fin membrane at level of ninth and tenth spines; proximal half of soft dorsal fin with numerous patches of orange pigment. Anal fin with orange pigment in an area from third spine through first and second softrays. Left pectoral fin with two oblique bars of orange pigment be- ginning near dorsal margin, slanting backward, and ceasing at about middle rays of fin; these bars crossing about middle of each ray involved; a small blotch of orange pigment near base of fin covering middle rays. Pelvic fins with areas of orange pigment over soft- rays three through five. Remainder of specimen pale except for some dusky pigment on distal part of posterior portion of spiny dorsal fin. A specimen (41 mm SL) of Scorpaenodes tredecimspinosus was obtained east southeast of Cape Fear, North Carolina (33°37.4’N, 77°07.5’'W, depth 37 to 38 m, R/V Eastwaprb sta. 23151, field no. GMBL 73-196, 2040-2112 hrs, 8 No- vember 1973, by trawl). This species is distributed widely in the tropical western Atlantic—localities of capture include Florida, the Bahamas, Dominica, Hon- duras, Panama, Venezuela, Dutch West Indies, and Los Roques in depths of about 8 to 82 m (Eschmeyer, 1969). The collection of the specimen off Cape Fear represents an extension of the known range of the species of approximately 580 km northeastward along the Atlantic coast of the United States from the most northerly locality (29°09’N, 80°12’W, R/V SiLver Bay sta. 4419) listed by Eschmeyer (1969). A specimen (60 mm SL) of Ectreposebastes imus was collected southeast by east of Charleston Light, South Carolina (32°11’N, 78°56’W, depth 256 m, R/V Orecon II sta. 11732, field no. GMBL 72-42, 1815-1930 hrs, 23 January 1972, by trawl). This species has a very extensive range in tropical and sub- tropical waters, having been reported from the eastern Pacific, off Hawaii, off Japan, eastern Atlantic, and western Atlantic—off the Mississippi Delta, Hon- duras, Colombia, Puerto Rico, and northeast Florida (Eschmeyer and Collette, 1966; Eschmeyer, 1969; Maruyama, 1970; Collette and Uyeno, 1972; and Esch- meyer and Randall, in press). Although Eschmeyer and Randall note that E. imus No. 3, 1975] ANDERSON ET AL.—SCORPAENID FISHES 173 is a near-bottom species, some individuals have been obtained by midwater trawls at considerable distances off the bottom. The capture of the specimen reported herein represents an extension of the known range of the species of ap- proximately 280 km northeastward along the Atlantic Coast of the United States from northeast Florida (29°54.5’N, 80°10’W, R/V OrREcoNn stas. 5233 and 5234). An abnormal specimen (54 mm SL) of Scorpaena agassizi was collected south- east by south of Cape Lookout, North Carolina, in the same tow as the Pontinus nematophthalmus discussed above. The usual number of dorsal-fin spines in this species is 12, but our specimen has only eight. A radiograph shows that the spiny dorsal fin and anterior pterygiophores are grossly distorted. The first dorsal-fin spine is quite small and far anterior to the rest of the fin. The second and third dorsal-fin spines are absent, as is the antepenultimate spine even though the pterygiophores with which it would be associated (if it were present) appear normal. We have not been able to determine which spine is the fourth one miss- ing from the usual dorsal complement. In other respects this specimen agrees with the description of S. agassizi as given by Eschmeyer (1965). Eschmeyer (1965) noted a similar abnormality in Scorpaena isthmensis. We thank Gene R. Huntsman (National Marine Fisheries Service, NMFS) for inviting two of us to accompany his group on cruises of the R/V Eastwarp and Bennie A. Rohr (NMFS) for aiding our collecting activities aboard the Orecon II. William N. Eschmeyer (California Academy of Sciences) sent us a copy of the pertinent pages of a manuscript that he and John E. Randall have in press. Norman A. Chamberlain (Grice Marine Biological Laboratory, GMBL), Bruce B. Collette (NMFS), and William N. Eschmeyer reviewed the manuscript. The material examined is housed in the collections of the Grice Marine Biologi- cal Laboratory. This is contribution number 35 of the Grice Marine Biological Laboratory, College of Charleston. LITERATURE CITED Co.tetteE, B. B., anp T. Uyeno. 1972. Pontinus niger, a synonym of the scorpionfish Ectreposebastes imus, with extension of its range to Japan. Japanese J. Ichthyol. 19:26-28. EscuMEYER, W. N. 1965. Western Atlantic scorpionfishes of the genus Scorpaena, including four new species. Bull. Mar. Sci. 15:84-164. . 1969. A systematic review of the scorpionfishes of the Atlantic Ocean (Pisces:Scorpaeni- dae). Occ. Pap. Calif. Acad. Sci. 79:i-iv + 1-143. AND B. B. Couette. 1966. The scorpionfish subfamily Setarchinae, including the genus Ectreposebastes. Bull. Mar. Sci. 16:349-375. AND J. E. Ranpa.t. In press. The scorpaenid fishes of the Hawaiian Islands, including new species and new records. Proc. Calif. Acad. Sci. Maruyama, K. 1970. Some deep-water fishes from off the Tohoku and adjacent regions. Bull. To- hoku Reg. Fish. Res. Lab. 30:43-66. (In Japanese, not seen in translation.) Florida Sci. 38(3):171-173. 1975. Biological Sciences NOTES ON THE INTRODUCED GECKO HEMIDACTYLUS GARNOTI IN SOUTH FLORIDA RoBERT Voss 7393 SW 79th Court, Miami, Florida 33143 OccuRRENCE of breeding populations of the gecko Hemidactylus garnoti in south Florida was first noted by King and Krakauer (1966) who reported that it had been introduced prior to 1964, and existed in two widely separated locali- ties in Coconut Grove and southwest Miami. Truitt and Ober (1971) subse- quently also listed it as occurring in the area. For the past 7 or 8 years I have observed this lizard in and about my home in southwest Miami. Since 1966, the range of H. garnoti has spread, and it is now found through- out most south Miami suburbs. Where found, it is relatively common, strictly nocturnal, and measures 50-60mm (snout-vent). At night, the adult is a uniform ash-grey save for the orange ventral surface of the tail. When discovered in hiding during the day, H. garnoti is a darker brownish-grey with lighter mottlings. The newborn young are darker than adults and have lightly banded tails. Hemidactylus garnoti is most frequently seen on warm nights on lighted walls and on the screens of lighted rooms where it feeds upon the moths attracted there. It actively pursues its prey in a series of short, quick rushes. During the day I have found H. garnoti under dead leaves and rocks, and under the bark of trees. Like H. turcicus, it squeaks faintly when caught. I have not heard it vocal- ize on other occasions. The eggs are nearly round, smooth, white and brittle, measuring 7-10mm in diam. They are laid, invariably in pairs, from June to January in enclosed dry places such as flower pots, gardening gloves, and piles of brick. They are often cemented together, or to some hard surface. After an incubation period of about 60 days, the young hatch. The newborn lizards have a snout-vent length of 24- 26mm. Despite occasional cold winters, H. garnoti appears to be doing well in south Florida and its range appears to be expanding. As we have no native nocturnal lizards in the area, H. garnoti fills a previously unoccupied niche in which it may be expected to continue to thrive. Thanks are expressed to Dr. Roger Conant who encouraged publication of these observations. LITERATURE CITED Kinc, W., anp T. Krakauer. 1966. The exotic herpetofauna of southeast Florida. Quart. J. Florida Academy Sci. 29:144-154. Truitt, J. C., anp L. D. Oper. 1971. A Guide to the Lizards of South Florida. Hurricane Press. Miami. Florida Sci. 38(3): 174. 1975. Biological Sciences KEY TO THE MOSSES OF PUERTO RICO Harvey A. MILLER AND KeiruH W. RussELL Department of Biological Sciences, Florida Technological University, Orlando, Florida 32816; and Science Library, University of Houston, Houston, Texas 77004 Asstract: Identification of genera included in Crum and Steere Mosses of Porto Rico is simpli- fied by provision of a generic key not included in the original work. The key is artificial and based upon vegetative characters. Stupy of mosses of Puerto Rico in connection with ecological studies of an elfin forest organized by Richard A. Howard (1968) led us to recognize the need for a key to genera of mosses. Crum and Steere (1957) did not provide a key to larger groups in their manual MossEs or Porto Rico AND THE VIRGIN IsLANDS thereby limiting its maximum usefulness to those already pretty well grounded in bryology. Accordingly, we undertook preparation of a key to genera as repre- sented by the 268 taxa treated in the manual. Further, we sought to provide for identification of sterile material with the inevitable result that some genera occur more than once in the key where a contrasting pair of characteristics might be represented by different species within a genus. The key is totally artificial and limited to the flora noted. It cannot be reliably used elsewhere—indeed, the Puerto Rican species is sometimes unusual within the genus (e.g., Tortula) and could lead to serious mis-identifications in other floras. We hope that the ex- istence of the key will stimulate collecting by students and others resident in Puerto Rico so that we may learn more of its most interesting bryoflora. We thank Richard A. Howard for the opportunity to participate in the elfin forest study supported by a National Science Foundation grant (GB-3975) to the Arnold Arboretum. Howard A. Crum most kindly reviewed an early draft of the key and provided valuable suggestions. KEY TO GENERA The number which precedes the generic name is the page number where the key to species or generic description will be found in Crum and Steere. 1 Branches in fascicles, leaves dimorphous on stems and branches, large empty porose cells in a network of elongate, narrow chlorophyllose cells ....404. SpHacNUM 1 Branches dichotomous or pinnate but not regularly fascicled, leaves rarely di- morphous, leaf cells essentially homogeneous in upper blade .....................0000e 2 2 Plants erect and unbranched or dichotomously so, female sexual buds terminal on mainstem’ (AaCrOCarpOUS MOSSES) ........2...4..c0c.siesess seen este eee teeee eee ees 3 2 Plants prostrate and usually freely branched, female sexual buds lateral and several on a single stem (pleurocarpous mosses) ...............:.0:0:00ceee 57 176 UE 11 13 13 15 15 1 17 19 19 FLORIDA SCIENTIST [Vol. 38 Leaves with numerous lamellae on upper surface of the costa, peristome with a diaphragm and 32 or 64 teeth (Polytrichales).....................0:.:c::2:-:-210s520 4 Leaves without lamellae on upper surface of the costa, peristome toothed or absent, rarely diaphragmed (Bryales) ...0....0....:+:-2-.t.s:40% @eue-¢-ete- tate eee 5 4 Leaves 3—3.5 mm long, bordered by elongate cells ................ 587. ATRICHUM 4 Leaves 5—7 mm long, not bordered ...............ceeeeeeeeeeeees 584. PocONATUM Leaves in two ranks at the insertion, the antical base of the leaf split into con- duplicate blades .....2:ss.c.-ctise8 snes stoiscess sees ee 407. FissIDENS Leaves in three or more ranks, more or less transversely inserted, leaf blade single throughout .......:..00 Line Mn eer 6 6 Leaves whitish, mostly several cells thick with outer layers of enlarged leucocysts with a single+median layer of reduced chlorocysts, leaf blade reduced and costa comprising most of the leaf (Leucobryaceae).......... if 6 Leaves green to blackish or brown, blades unistratose, costa lacking leu- COCYSUS....1..cc.cceesseeisnesovetersaronstnngens seentenee sinew chee ei rr 9 Leaves with a false costa (stereid band), often toothed on back or with serrulate MAT PINS 6/5 lie. Ee Ne REE OP CO cl eee ee 444, LEUCOPHANES Costa homogeneous throughout, margin of the costa entire ......0.0.00..0 eee 8 8 Leaves plane, oblong above with parallel sides abruptly narrowed to cuspidate tips... eh ee eo ae ene eee 444, OCTOBLEPHARUM 8 Leaves grooved, tapered above, tip acute to acicular.......... 441. LEucoBRYUM Inner basa! cells much enlarged and hyaline, sharply differentiated from the blade (Calymperaceae) oo.) 0 iic ee dete teteeccse i cent teed ee 10 Inner basal cells various, but not hyaline or sharply differentiated ............0....0..0.... 11 10 Leaves almost always with narrow intramarginal bands of transparent linear cells 2—12 cells from margin, sometimes extending only a short distance above the base... .5,.cct este ee 446. CALYMPERES 10 Leaves lacking intramarginal bands of elongate cells, unbordered, or bordered at margins only with pale elongate cells, or with serrate WANS ure Le IN casa tata, SREB, ee ee 452. SyRRHOPODON Leaves ecostate or costa very short, sometimes double ..............0..0.0cee ee 12 Leaves costate, costa single. 3....4.:c-.seenssee.ees ste. dee 13 12 Leaves narrowly lanceolate, margin serrulate to toothed, plants minute (Micromitnum, see Crosby, 1968) =... 2 ee ee 484. NANOMITRIUM 12 Leaves ovate, margins entire, plants medium to large ............ 535. HOOKERIA Leaf cells smooth or papillose only by projecting ends of elongate cells, cells mostly elongate to rhomboid above .......:.:..0...0..)...0) lon... ee 14 Leaf cells pluripapillate to rounded or conic papillate, cells mostly isodiametric DDOVE w.cccecesin iol sshd Sandu op ies artes chivneduadeodssetiauan obeerenahe cage Heed oe 36 14 Alar cells sharply differentiated, often pigmented and auriculate................ 15 14 Alar cells undifferentiated from other basal cells .............. 2. 22a 17 Leaves with a hyaline border of elongate cells «0.0.0.0... 439. LEUCOLOMA Leaves unbordered or border faint and not hyaline ...........%....).........2-.34) eee 16 16 Costa narrow and about the same width over its length, leaves strongly toothed above the expanded base... +)... eee 438. HOLOMITRIUM 16 Costa broad below, tapering to the apex, leaves entire or denticulate near the tip, leaf base auriculate in the alar region but otherwise not ex- panded! 1. 202) 1 UT) SOO Tae ea aan ee eee 433. CAMPYLOPUS Branching dichotomous or none, rarely in a comal whorl, stems leafy through- out, leafless prostrate primary stem! absent ...)0...0...0.00..20). 2.2.2 18 Branches erect and abundant from a leafless prostrate primary stem ...................... ey) 18 Leaf cells elongate, thick-walled and papillose by projecting ends.............. 19 18 Leat cells variously shaped but smooth =...) 224) 20 Leaves plicate at base; basal cells linear... = eee 493. BREUTELIA Leaves not plicate at base, basal cells rectangular to isodiametric .. 494. PHILONoTIS No. 3, 1975] MILLER AND RUSSELL—MOSSES OF PUERTO RICO 177 20 Leaf cells rounded-quadrate above, at least as wide as long, leaves un- JORIGTEL, ccosscudadeieejanleane mses seeks ett atte nine Rice eA Nee suai ee ee 21 20 Leaf cells elongate rectangular to rhomboid, leaves sometimes bordered eeminaneneixclomeatercell Spates 8. ei. wisn utter lee Lie rst Maeda hates oon sD done 23 21 Leaf margin thickened and doubly serrate |... 492. RHIZOGONIUM 21 Leaf margin one cell thick and entire or singly toothed «00.00.0000. eee 22 aaeiteavesiobovate, sheathingvat the base 0.2).....00 i000. 463. RHAMPHIDIUM DET ASEN ES JOD CS EEG O00 0 oP aR oS ee eRe ee 474, BARBULA 23 Costa subpercurrent to excurrent, leaf blade often narrowed above and sub- TLE TE scondessanceuebacedoesss pes NOgUR a dae NODS e ae Anema Eo Pe eR ack As ee a 24 23 Costa ending well below the leaf apex, leaf blade broad above and often TI UIRG SESE UTS ON OES cask Ne 2 aeRO Ae Cai Sg ee ee er ee ee et 28 24 Leaves with an expanded base and a narrowed tip, often subulate and comprised entirely of a strong excurrent Costa .o.00.0..0.. cece 25 24 Leaves ovate to elliptical or obovate, widest above the base, costa per- SORE MMU RS MONE CX CUTMEMNG ies ne ce ss ceca ie conte ces ero osc ty ee owess cdveete ntact nseusebcesnsvenetees 32 25 Leaf blade oblong, narrow above and extending around the subpercurrent ERSESEEMETOMEUSE APEXTOOLNEE 21k ccs. c ise sen ene vases ov enta se sbeysees ...... 430. TREMATODON 25 Leaf blade tapering toward the acute to acuminate tip .......0000.00. cee 26 26 Subulate leaf tip about 3 times longer than leaf base, seta strongly CULE Cs enqonc laaieea cada oe tae Nene dinate knit a eee 430. CAMPYLOPODIUM 26 Subulate leaf tip less than 2.5 times the length of the leaf base which RaplessuOrtip .Seta CLeCt aNd StraiSlat (ess..05.e sess. cee gs seyenset ae ie seosuae server ented wcobosoase 27 27 Leaves strongly clasping, capsule stomatose, peristome 250—450 p high.......... ATARI ated secs esate ddcuntnshsebeanea Wiabevunavovvetes 424. ANISOTHECIUM 27 Leaves not conspicuously clasping, capsule without stomata, peristome rarely 22 PLD BS SON Ore ee ee eee 425. DICRANELLA 28 Leaves dimorphic, lateral leaves elliptical and dorsal leaves smaller and lanceolate, cells lax, becoming very narrow along the reddish bordev.... sccecceesltbay salad i Oat Soli SMe nU Ban ARG ee ner a ee aa ee 486. EpiIpTERYGIUM 28 Leaves all alike, cells lax with unbordered leaves or, if leaves are |SCIG SEG Ld OST ACCS) UGH nih 00 Weare ge alee tennn een en ert een Meee eet e, 29 29 Leaf cells lax, thin-walled, rectangular-rhomboid becoming much enlarged pa NPE PRC SHUNT ORCL T CCld 5 are 8 eh SS oo cso os tec vance a suetnidanbcniec onc idive ROMO: 30 29 Leaf cells firm, rhomboid-hexagonal above, elongate below, leaves distinctly wearmmetcammtn anand of elomgatelcells) x25.fcs5.5.2 42. ctaicdec serene caceecsee supigbedesaeote neck 31 30 Costa ending well below apex, leaves 1.5—2 mm long ...... 431. WILSONIELLA 30 Costa usually ending 2—5 cells below apex, leaves 0.3—1.3 mm long...... sensed: ite loeetaphdbe gee eeO ae MRPONS Sedo ane een: 479. SPLACHNOBRYUM 31 Border 1—3 cells wide all around, leaves not keeled.......................... 534. LESKEODON 31 Border 5—15 cells wide at base, 2 cells wide at apex, leaves keeled with a "TEAST T8) Cuesta che SBN IEIE ROLE AUR. APE MERRIE IS tee Sache Mee ee 532. DALTONIA 32 Leafy shoots arising from a prostrate, rhizome-like stem, leaves ap- pearing whorled above and more than 10 mm long .......... 487. RHODOBRYUM 32 Leafy shoots erect, leaves all along stem, though sometimes larger above, RI EOSS UEAUL SEMIN LOIS a8 beste athe eed Shen Rents ges MA LA essen ne 33 33 Leaves plane or nearly so, leaf cells lax and thin-walled, rectangular to ob- MSEC IE OLCCM ANIC! (JAFETICHY MAL OUS 02 Jost 2246005 eh cise tet east acd seek Rectan 34 33 Leaves mostly keeled, leaf cells firm, short-rhomboid to narrowly rectangu- RPmSSCAMEMSILANLY ICME MILE CATO IEEIN 3520.0. 25 -.npoddnen aeadayniees smecbeatinaton, losses Qusecesedlnn-- 35 34 Leaf margin entire or weakly toothed above, costa subperrent to short- GIG ISTE cess Sects RRC Ee LET eR eee 483. FUNARIA 34 Leaf margin denticulate in upper half, costa ending well below apex...... _aenunaniceeapartb bebe th weed Sha SSC Oo ROR ET eee eS ee EEE NOR reer 483. ENTOSTHODON 178 35 35 37 37 39 39 4] 4] ol ol FLORIDA SCIENTIST [Vol. 38 Stems bearing large reddish propagula, leaf cells oblong-hexagonal, up to 3 times.as long as broad): ..1.1.4..::2s024.qi-eee enna oe 486. BRACHYMENIUM Stems lacking propagula, leaf cells usually more than 3 times as long as brad wis sccsceuassscecnesencdendeawnnsesdceonevedinasnee ce seuseone ss 0taS Ae a rr 488. Bryum 36 Leaves distinctly bordered by a band of elongate thick-walled cells, alar cells inflated or strongly differentiated................0..0...... 439. LEUCOLOMA 36 Leaves unbordered above or indistinctly so, alar cells undifferenti- ALO oo. eiccecsecseseeteetsndececeos snes teceecnnneceetnoes vanesn 1 etlel soe = 37 Branching dichotomous or none, stems leafy throughout, leafless prostrate primary stem absent .....2....0...0.c..ccecdconsessesseseessawsetneew soos 555 38 Branches erect and abundant from a prostrate leafless primary stem...................... 53 38 Leaf cells thin-walled, elongate to rhomboid below the apex, basal cells thin-walled and lax |... saci: Mencieen ee 39 38 Leaf cells mostly thick-walled and isodiametric, basal cells often some- what elongate and usually thick-walled... eee etter 40 Leaf apex broadly rounded on spathulate leaf, costa ending well below apex.... Te rr et Re eee eth PMs ME erg he SanccSaataronsconsen: 481. GYMNOSTOMIELLA Leaf apex obtuse on ovate-lanceolate leaf, costa subpercurrent ..463. RHAMPHIDIUM 40 Leaves with a band of elongate, hyaline thin-walled cells extending up the margin from. the leaf base ........,...........j:.ss:sseseeesee++9985t | er 4] 40 Leaves with cells of basal margin undifferentiated or at least not hya- VEIN so ons cgeceiede $s sateen ndsgocbatc node oudeauenaeatie let vandop deste tyre eceeaee aoe eee eee rr 42 Costa ceasing shortly below leaf apex, upper cells not papillose, leaves 2—2.0- mm-long; keeled: o.)-si7 sets oes sata actos come eee 479. LuISIERELLA Costa excurrent, often as a mucro, upper cells densely papillose, leaves 3—9 mm LOWS. oysecs uk evstasoovt cde Hoek sear tlou Mopcehiiso tae ae vetcbe. eee ne oe cee ee ee 470. TORTELLA 42 Leaf margins plane when wet, sometimes slightly involute near the CD sos Sesh nee nc ceeneetn tee cagilnnn Osan taetourom uate Roe et aes 43 42 Leaf margins curled, either involute or revolute near the base .................... 46 Leaves oblong to lanceolate, gradually tapering to the base ....................eee 44 Leaves long and slender with a somewhat expanded leaf base broader than the Dae aise a fehseed ne eahpeei on Seu onde He osenes eased onde nee ne sae se aacegies a eee ee rr 45 44 Stems radiculose, leaves keeled ..............000cccccceeeeeeeeeeeeeeees 459. ANOECTANGIUM 44 Stems not radiculose, leaves not keeled ............00........cceeeeeeeeee es 473. HyopuHILa Costa excurrent as a hyaline or yellow mucro, leaves usually involute at mar- gins, at least: abOve 30: bs es. se: 15, aver ecpeaenehee a eee eee 466. TRICHOSTOMUM Costa subpercurrent,.margins plane)./ 2272. eee ee 464. TUERCKHEIMIA 46 Leaf margins involute throughout or at the base 2.0.0.0... eects 47 46 Leaf margins revolute throughout or at the base ...0..0.....00 ccc 49 Leaves 1.5—2.0 mm long, leaf tapering to a lanceolate tip ...... 460. HymMENOSTOMUM Leaves 2.0—3.5 mm long above (rarely 1.5 mm below), leaves oblong above from an expanded base .2......0:...5 0). c:eededeedeceitesnsstoeteiastas cided lee err 48 48 Leaves sharply keeled above ...2..0:..c00:eseens ee ee 461. WEIssIA 48 Leaves involute but not sharply keeled ........0.....0...00.0... 466. TRICHOSTOMUM Leaves bordered by a band of elongate cells 000.0000... cece 478. ToRTULA Leaves unbordereéd iii. :cceseceteliiedicl dbessostedielgess dada eee ee 50 50 Leaf margins denticulate to dentate above, papillae C-shaped.............. ubddiodd tulle ioOA NG dan DCRR NE Rr era es 476. BRYOERYTHROPHYLLUM 50 Leaf margins entire, papillae various «00.00.0000... cesses eeeeteeteeteeeetenees ol Leaves widest at about the middle, plants small and bud-like, upper leaf cells thin-walled, capsule immersed .......:....::4....40:.4120- 477. PHASCUM Leaves widest near the base, plants with elongate stem, upper leaf cells in- crassate, capsule exserted. .......22:0...:.0.seteeisneurteeeneesstnete ee 52 02 Costa disappearing below the apex ................cee 462. GyYMNOSTOMUM o2 Costa percurrent to excurrent .4.5...-+.0.32.--..--a ee 474. BARBULA No. 53 53 eye) 59 o7 o7 59 59 61 61 63 63 65 65 67 67 69 69 71 71 3, 1975] MILLER AND RUSSELL—MOSSES OF PUERTO RICO 179 Leaves bordered at the base by elongate cells, inner basal cells short _ oa sop ogeageeote hah Von AGpeascitE Nac lcd RRM ct cnet AO A EE 498. GROUTIELLA Leaves unbordered all around, basal cells all elongate ...00..0.000 00: 54 54 Secondary stems not tomentose, leaves usually 2—6 mm long............ ncccannscechendsocen cA GLEEg SAE Se Eee pce nee 5900. MACROMITRIUM 54 Secondary stems tomentose, leaves usually 1.5—2.0 mm long.............. _ opcingheestt < Sgansacees A SARAce che AAaMing nee Ran ns cs Peat ne ae eee 905. SCHLOTHEIMIA Leaves bordered at the base by elongate cells, inner basal cells short... _ 222 202 92 og8 Stee ed Ra cRoacaeck aspera eey a pe OO ae ee 498. GROUTIELLA Leaves unbordered all around, basal cells all elongate ....0..00..00 0 occ 56 56 Secondary stems not tomentose, leaves usually 2—6 mm long........... RS Re: SEO iene ene ets eA RAR 900. MACROMITRIUM 56 Secondary stems tomentose, leaves usually 1.5—2.0 mm long............_. Beare PN ON by Peis Ae PALES ehh DUS Aah Mon aba Pay 905. SCHLOTHEIMIA ere auc Ost ausingl essere ere her A ECR DHEA RA danas 58 Me mresnmel costa double OrdlOnen yf. Fie ced ee ANN tele eech hiew 86 58 Leaves dimorphous, lateral leaves large and complanate, dorsal leaves smallerand nearly transversely inserted 2.00... c..iececeseccaceecersnsenecrseneracnesne 59 58 Leaves all of one type and of about uniform size oo... eee 61 Leaves strongly inrolled from tip when dry..........0.00000 cc. 5906. HELICOPHYLLUM Leaves somewhat contorted but not inrolled when dry ..........0...0.. cece 60 60 Lateral branch leaves bordered, costa ending about 3/4 leaf length... -c2 sss sened8686SepomeBetaae SO USBBA aren Bete -e Nara c nt a fiets Nas nS noth eileen ie 904. HyPOPTERYGIUM 60 Leaves not bordered, costa excurrent .........0.00.0000ccccceeeeeeeeees 507. RHACOPILUM (ZELL EE_F DECI OSS nc opcacsanaseaagenslioe oe asuB aPON CRG REA oe MENA tea anes neni ace iter 62 we BS SELLE SLO OTD coc csc ossaeraetage ee less a eed RUSS aR ec cP ROE 69 62 Leaf cells isodiametric or short and up to 3 times longer than broad .......... 63 62 Leaf cells elongate to linear, at least 5 times longer than broad .................. 65 Paraphyllia absent, leaves 1.5—2.0 mm long ..............0.0..c 064. STEREOPHYLLUM Paraphyllia present, usually abundant, leaves less than 1.2 mm long ...................... 64 64 Apical cells of branch leaves bearing single sharp papilla.................... _ Svc nscale de see ede bcoO eee Bete SOE OIC, ane SE ase aI 508. HAPLOCLADIUM 64 Apical cells of branch leaves bearing 2 or more papillae ........ 596. THUIDIUM Leafy stem complanate with flattened branches ...............0..0....... 527. POROTRICHUM Leafy stem not flattened, with leaves equally distributed around the stem ............ 66 66 Leaves acute to acuminate, upper margins plane ........ 511. LEUCODONTOPSIS 66 Leaves attenuated to a slender hairpoint, upper leaf margin often un- ID ELS, ‘2. ede sacbstcl die tieadtie ae caters «el ema an ar a am Cn ne RONMOR dee SOE 67 “ELE UNTER TiN ae aa ecg I ee ne NO eee ne 519. PAPILLARIA 2a 5 TWAT ASS AE core ae ge ete Be re Me IN pee a nie INC tare ey net ely RPP Sales een 68 68 Branches terete, densely foliate, branch leaves abruptly acuminate EMME POUNCE MALT POMC. n ccc. cc2 fou ops coetraren waned snennnynee nt 520. METEORIUM 68 Branches slightly flattened, laxly foliate, branch leaves gradually acu- minate forming a filiform, flexuose-crispate point ............ 521. AEROBRYOPSIS Leaf cells above isodiametric to short-ovate or rhomboid, up to 3 times longer Se PMTE AY A0 ee ck hey foc cgsncuiioesddeaneacaucapgbaurore Sonceegs ens 70 Peat cells linear, atleast o times longer than broad .z.............::5::0-.:++.0+sse.-ceeeesee- >= rit 70 Leaf margins distinctly bordered by a band of elongate cells ...................... 71 RO Mia ATEN OTS URN OTC oo eee sees rap tots ces dno ess yanndvrevngntatonspeonse veri (e Border 1—3 cells wide all around, leaves not keeled.......................... 534. LESKEODON Border 5—15 cells wide at base, 2 cells wide at apex, leaves keeled with a TLS LIALT TUG snereiscsae ccataengh deste eaeeheg Social” ON EP eer mentee nent nan 533. DaLTONIA 72 Leaf gradually tapered to an acute to slenderly acuminate tip .................... 73 72 Leaf broadly rounded to truncate or obtuse at the tip ...............0.. 75 180 85 85 87 89 91 93 FLORIDA SCIENTIST [Vol. 38 Leaf margin somewhat inrolled, entire to minutely serrulate above.............. si Rewauhiogne «ADM ka Sek ooh glk cntoe nasi Raeaean ane en eer 509. ACROCRYPHAEA Leaf margin plane, crenulate by bulging cells .:.........)...3..20....ssussress eee 74 74 Costa extending 2/3 to 3/4 of leaf, margins crenulate toward tip, cells rhomboid: 2s...s:ce.eses aie. dhies- ante ee ee 595. HELICODONTIUM 74 Costa extending 1/2 to 2/3 of leaf, margins entire or subserrulate at branch tips, cells oblong-linear 2-001. 500. FABRONIA Leaves ovate becoming narrower toward the apex..................05. 527. POROTRICHUM Leaves oblong, about the same width to near the truncate apex..............0.... 76 76 Leaves truncate to emarginate at apex ........... ee 524, NECKEROPSIS 76 Leaves broadly rounded and toothed at apex ................. 525. HOMALIA Leaf margin entire or serrate in the upper half.......................).. 78 Leaf margin strongly toothed nearly to the base... :22.:....2.000.)..s-a ee 96 78 Leaves plicate below ..........:..:.0..c:stecne nn eunennetes ie 79 78 Leaves plane to concave but not plicate below 2.0.0.0... eee 80 Leaves 1.5—2.0 mm long, costa ending at, or slightly above, mid-leaf............ IE ee er ee RI em SPREE TAT ban, bE e Moho dL Abechincodnoaase: 561. BRACHYTHECIUM Leaves 3—4 mm long, costa ending in acumen .................... 509. DENDROPOGONELLA 80 Upper leaf margins inrolled below, abruptly acuminate tip borne on a broadly:oval blade: tc. sti. ctot0t.3 232 ein cipes cameos See eee 516. SQUAMIDIUM 80 Upper leaf margins plane, leaf tip acute to acuminate from ovate or Oblong blade .......i.5.....isceness+-nseceessosneeostsbeseaebeopearecuale eaeeane err 81 Alar cells short, rounded to quadrate, sometimes yellow «0.0.0.0... eee 82 Alar cells undifferentiated .......:.....:c0..0:c..se-aceescavneentsieescneaeedoapeieuce pee 85 82. Costa percurrent or nearly So .........0:.:0:4..ton he 83 82: Costa ending near mid-leaf -............:1......0ecceeesreesotn cous se sneebe pegs oe 84 Leaf axils usually bearing filiform microphyllous branchlets, leaves imbricate When diy co... ses Ssactenn dene eels nate teat ee eee 512. PsEUDOCRYPHAEA Leaf axils without microphyllous branchlets, leaves erect spreading ..514. PrREELLA 84. Alar cells poorly differentiated). .......5.402.s0.50.2-1s4/..822 nonsense 514. PIREELLA 84 Alar cells well ditterentiated: ...-2-.2-7 2 eee eee 964. STEREOPHYLLUM Leaf margins emtine Ao54 cies. cca0 esans-soneetctee. cee, eee ee ee 508. CRYPHAEA Leaf margins serrate or serrulate, at least in upper 1/3 ................ 527. POROTRICHUM 86 Leaves in two ranks (distichous) at insertion on the stem, or leaves di- morphous with dorsal leaves erect and reduced ...............0::::cccseeeeerrteeee 87 86 Leaves inserted all around the stem and of one type, although some may be complanate but not distichous or dimorphous ..................::::cceteterees 88 Leaf cells densely papillose, leaves less than 1 mm long .................... 498. ERPODIUM Leaf cells smooth, leaves 2—4 mm long «0.0.0.0... cece: 523. PHyLLOGONIUM 88 Costa extending to beyond mid-leaf .....0............0...50000004.00-2 ee 89 88 Costa short, rarely attaining mid-leaf, or none «0.0.0.0... cc eceeeeceeeeteteeeteees 98 Leavesstrongly plicates 0 i..0c26fcccicseoce ese ee 502. HEMIRAGIS Leaves plane to concave but not plicate ...................2.00c-5eeqeer 242-2: 90 90 Leaf cells thick-walled, isodiametric to oval, undifferentiated or slightly longer atthe base .....52......c:cc0ccsnescessngsees:songenresesoestes Rage seer 91 90 Leaf cells thin-walled to firm, cells rhomboid to linear becoming lax below occ il esi aesiath cane sedis dg ee 92 Costae ending well below the leaf apex, without dorsal spines .... 530. PrLorricHum Costae subpercurrent, ending abruptly in blunt dorsal spines.. 530. PiLorricHip1uM 92 Leaf cells nearly isodiametric or laxly hexagonal .................0.:::ccceeeeteees 93 92 Leaf cells narrow to nearly linear and prosenchymatous .................:.:0000+ 94 Leaves bordered, leaf cells smooth .........00.......0..0.000cccocececeeeceeeeseees 536. CyCLODICTYON Leaves unbordered, leaf cells usually papillose....................0..... 538. CALLICOSTELLA 94 Leaf cells bearing 3 or 4 papillae 00.0.0... ccc 548. HyYPNELLA 101 101 103 103 105 105 107 107 109 109 Ill lll 113 . 3, 1975] MILLER AND RUSSELL—MOSSES OF PUERTO RICO 181 PE LGR GEIS RINGGUN seocgecanaacesee 29° 0e bee cbe ace cEer oe EE Hap ITE Gee ene ae ae oe 95 Lateral leaves asymmetric, leaf margins plane ..............0...0..000. 539. HooxkeEriopsis Leaves uniform, leaf margins recurved, at least in lower 1/2 ..543. AcTINODONTIUM LE CON TEODOR ON oer beeline Saeco ete ee era ee eee ee 514. PIREELLA Sen Costaiendine wellibelow the tip ce. - 5.1025. oe-chesspse-tudenseeeveesandsoeteneetens teasvaaeeeseende 97 Leaves wide-spreading from the stem, leaves more or less plane 522. METEORIOPSIS Leaves erect-spreading from the stem, leaves concave .......... 562. RHYNCHOSTEGIUM 98 Leaf cells papillate over the lumen or lateral walls, often pluripapillate .... 99 98 Leaf cells smooth or papillate only by projecting ends of cells .................. 102 AAT GELS WOOT eieeIN Te. AE rs cacme serie dec Renee or cner aera ee rae tev eee en 100 Alar cells differentiated with 2 or 3 often inflated and sometimes pigmented ...... 101 100 Leaves concave, oblong-ovate, papillae several, multifid ...... 548. HyPNELLA 100 Leaves complanate, broadly lingulate, small papillae at the upper cell ends and | or 2 near the middle of the cell .......0.0.00.0.... 576. GLOSSADELPHUS Leaf cells unipapillate, leaves with long flexuous tips .............. 074. TRICHOSTELEUM Leaf cells pluripapillose, leaves acute to acuminate...................... 575. TAXITHELIUM 102 Leaf cells laxly rhomboid to elongate above but not linear or nar- oma aS ITU OSC Repeat se ern ech ted seh ass. oyns. LUBE Reds ol ashame anpacheads 103 102 Leaf cells mostly firm, rarely thin-walled, and linear to sinuose ................ 112 Leaves with alar cells inflated at the basal angles.....0..0..0..0.0. cc ceeceseeeeereeeteeee 104 Leaves with alar cells not or slightly differentiated ...0..000.00.00 cece: 105 104 Upper cells rhomboidal, peristome single ..................0..0.... 968. MEIOTHECIUM 104 Upper cells elongate, peristome double........................ 568. SEMATOPHYLLUM Plants unbranched or nearly so, robust with ovate leaves 4—5 mm long, cells 48S OK UES ie ec ac Rae cree es on ee ee 535. HOOKERIA Plants branched pinnately to abundantly so, sometimes from a prostrate rhizome-like stem, leaves less than 4 mm long, leaf cells usually narrow .............. 106 106 Leaf base oval, deeply concave, abruptly prolonged into a long flexuose ASTITEY esas ado dzeth pe sb de: SoBSS MTeoR Oh EACERA | San ane a 549. STENODICTYON 106 Leave base various, gradually narrowed to the tip or blunt................0....... 107 Plants regularly pinnately branched, leaves grass green, falcate-secund.......... ie a Ee eh RO PR PP Saal ao sah 5. cals Suan data taribuiniacoe keeienait 581. VESICULARIA Plants irregularly branched or forming an expanded frond, leaves complanate or meee SLC AGNI OMUE TIVE IOWASH 23... cag. dese ede acat ones tndendeasedoersuses -azeiognactits sencende tances 108 108 Leaves oblong with a nearly truncate tip bearing coarse teeth.............. «sacs Cate are SOME 7 EAC OP ED BO RRND FE STRAT OAe SSC: aR EE OTe SRE RRO 520. HOMALIA 108 Leaves ovate or broader toward the base, often attenuate at the tip ........ 109 Leaves lanceolate, tapering to a long acuminate, coarsely serrate tip, branches TULITEG. BIER eee aie ae ea co ne ener eee 500. RHYNCHOSTEGIOPSIS Leaves broader with tip from a broader base, margin entire to serrate, branches MAROC ONAPIDIPISIrtes Patera 2s. te Pe LTRs AE Doss onidn Gbt they tasty veaston teu 110 110 Leaf margin entire, unbordered, ecostate, leaf cells 150—200 w........ sccesce salhnculee sek dé c# a GbE nc Rede {oD ee PONE TORE DE EH aer