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Albumin Evolution and Its Phylogenetic Implications in the Plethodontid Genera Pseudoeurycea and Author(s): Linda R. Maxson and David B. Wake Source: Herpetologica, Vol. 37, No. 2 (Jun., 1981), pp. 109-117 Published by: Allen Press on behalf of the Herpetologists' League Stable URL: http://www.jstor.org/stable/3891767 Accessed: 03-02-2016 22:17 UTC

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This content downloaded from 136.152.142.101 on Wed, 03 Feb 2016 22:17:42 UTC All use subject to JSTOR Terms and Conditions Herpetologica, 37(2), 1981, 109-117 ? 1981 by The Herpetologists' League, Inc.

ALBUMIN EVOLUTION AND ITS PHYLOGENETIC IMPLICATIONS IN THE PLETHODONTID SALAMANDER GENERA PSEUDOEURYCEA AND CHIROPTEROTRITON

LINDA R. MAXSON AND DAVID B. WAKE

ABSTRACT: The immunological technique of quantitative micro-complement fixation was used to study albumin evolution in the plethodontid salamander genera Pseudoeurycea and Chiropterotriton from Mexico and Central America. Antisera were prepared for representatives of the five species-groups of Pseudoeurycea and for the a and f3 groups of Chiropterotriton. A matrix of reciprocal tests was used to construct a dendrogram illustrating probable relationships. The a and p groups of Chiropterotriton are as distant from each other as they are from Pseu- doeurycea. The rex and gadovii groups of Pseudoeurycea are not well differentiated, and uni- directional tests to 13 additional species of Pseudoeurycea indicate that many species have been incorrectly assigned to groups. Pseudoeurycea cephalica is especially remote from all other species of Pseudoeurycea tested; it is even more remote from the species of Chiropterotriton tested. The cephalica, leprosa, and bellii groups are all well distinguished from each other and from the gadovii (including rex) group. Within Chiropterotriton p the bromeliacia and picadoi groups are sharply distinguished from each other. The great range of immunological distance measured in this study (from 0 to over 100) indicates that both of these genera are as differentiated genetically as is the North American genus Plethodon, and that the evolutionary history of each of these genera encompasses most of Cenozoic time. Key words: Albumin, Amphibia, Caudata, Chiropterotriton, Phylogeny, Pseudoeurycea

NEARLY hal-f of the living Of the eight genera currently recog- are presently included in one taxon-the nized in the supergenus Bolitoglossa, supergenus Bolitoglossa of the tribe Bo- Pseudoeurycea and Chiropterotriton are litoglossini, family . This the most generalized in morphology (see exclusively tropical group is found only review in Wake and Lynch, 1976). We in the New World; its only close relatives have chosen them for a detailed investi- are the genera Hydromantes and Batra- gation of albumin evolution, using the choseps, both of which are inhabitants of quantitative immunological technique of western North America (Hydromantes micro-complement fixation (MC'F). In occurs in Europe as well). The supergen- previous work we used this technique in us Bolitoglossa is considered to be the conjunction with starch-gel electropho- most derived lineage of salamanders, and resis and morphology to analyze salaman- has undergone an extensive adaptive ra- der phylogeny (Larson et al., 1981; Max- diation (Wake and Lynch, 1976). The re- son and Maxson, 1979; Maxson et al., lationships of its component lineages are 1979; Wake et al., 1978). Electrophoretic poorly understood, for there is no fossil and morphological analyses relevant to record and there has been extensive mor- the present study will be reported else- phological parallelism and convergence where. (Wake, 1966). Attempts are being made The 23 described species of Pseudoeu- to integrate diverse kinds of data into a rycea are placed in five species-groups, coherent picture of relationships; the four of which occur only in Mexico. The present report is the first to deal with fifth group is mainly Guatemalan, but two macromolecular evolution in the group. species enter Chiapas, Mexico. We se- 109

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TABLE 1.-Salamanders of the genus Pseudoeury- TABLE 2.-Salamanders of the genus Chiroptero- cea (after Wake and Lynch, 1976). triton (after Wake and Lynch, 1976).

Species Chiropterotriton a Chiropterotriton O group Species multidentatus species- bromeliacia species- bellii P. belliia,b group group cephalica P. cephalicaa,b P. conanti C. multiden- C. bromeli- P. cochranaeb P. galeanae tatusab aciaa,b P. werlerib P. scandensb C. arboreus C. cuchu- P. altamontanab C. magnipes matanusb gadovii P. gadoviib P. unguidentisb C. mosauri C. rabbib P. smithia,b P. melanomolgab C. xolocalcae C. megarhinus leprosa P. leprosaa,b P. robertsib P. anitae P. firscheini chiropterus species- picadoi species- P. juarezib P. mystax group group P. nigromacu- C. chiropterusb C. picadoi latab C. chondro- C. richardi stegab C. nasalisb rex P. reXb P. goebelib C. dimidiatusb C. barbouri P. brunnataa,b P. exspectatab C. lavae C. veraepacisb a Albumin purified and antisera prepared. 'Albumin samples extracted. priscus species-group C. priscusb

a Albumin purified and antisera prepared. b Albumin samples extracted. lected one member of each species-group for preparation of antisera (Table 1). Two groups of essentially generic level are MATERIALS AND METHODS recognized in Chiropterotriton (Lynch Plasma (preserved in phenoxyethanol) and Wake, 1978; Wake and Lynch, 1976). from 18 species of the genus Pseudoeu- The a group is restricted to the moun- rycea and 10 species of the genus Chi- tains of eastern Mexico, north of the Isth- ropterotriton was used as a source of al- mus of Tehuantepec. There are three bumin. Voucher specimens of all species species-groups and nine species in Chi- are in the Museum of Vertebrate Zoolo- ropterotriton a; we prepared antisera to gy, University of California, Berkeley. one species (Table 2). Chiropterotriton Albumin was purified from plasma of /l is mainly Central American in distri- five species of Pseudoeurycea (Table 1) bution; it occurs from just south of the and two species of Chiropterotriton (Ta- Isthmus of Tehuantepec to the Meseta ble 2). Due to the small size of the ani- Central of Costa Rica. There are two mals, plasma from several members of species-groups and 10 species; we pre- the same population was pooled. Purifi- pared antisera to one species (Table 2). cation was by single-step, polyacrylam- Because most species of Pseudoeurycea ide-gel electrophoresis (Maxson et al., and Chiropterotriton are small, rare, or 1979). Albumin was identified by its flu- both, it was impractical to produce more orescence in the presence of 8-anilino-1- antisera. However, the number of anti- napthalene sulfonate (Hartman and sera available was adequate to investi- Udenfriend, 1969). gate the relationships of the genera and Antisera to pure albumins were pre- the major species-groups. While we em- pared in male Dutch Belted rabbits by phasize those species for which tests of the following procedure. Rabbits re- reciprocity are presently possible, we ceived an initial intradermal injection of also report unidirectional tests for a num- Freund's complete adjuvant and albumin ber of other species belonging to the ma- (1.2:1). This was followed at 7 weeks by jor groups studied. an intradermal injection of Freund's in-

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TABLE 3.-Matrix of immunological distances among albumins of species of plethodontid salamanders of the genera Pseudoeurycea and Chiropterotriton.

Antisera Species tested B R C L S CB CM P. bellii (B) 0 39 74 52 43 84 88 P. brunnata (R) 53 0 72 48 15 67 77 P. cephalica (C) 78 49 0 64 62 90 107 P. leprosa (L) 72 53 88 0 35 92 88 P. smithi (S) 47 21 71 35 0 60 87 C. bromeliacia (CB) 79 53 90 92 71 0 104 C. multidentatus (CM) 88 66 100 57 87 98 0

complete adjuvant and albumin (1.2:1). min available. Despite the relatively low Three weeks later, 1 cm3 of albumin titers, the antisera exhibited standard be- solution was administered intravenously havior, with an average slope for all sev- in the marginal ear-vein; this was re- en antisera of 385, similar to slopes re- peated one week later. The rabbit was ported by other workers for mammali- bled from the marginal ear-vein or by an, anuran, and salamander albumins heart puncture one week after the sec- (Champion et al., 1974; Larson et al., ond intravenous injection. A total of 2-3 1981; Maxson et al., 1979). mg of albumin was administered per rab- MC'F tests were conducted in all pair- bit over the total period. Three rabbits wise combinations and the averages of were used for each immunogen, and in- these reciprocal tests are shown in Table dividual antisera were tested for purity 3. The percent standard deviation from by the criteria described by Wallace et al. reciprocity (Maxson and Wilson, 1975) (1973). Individual antisera were pooled was 1 1%, similar to that reported in other in inverse proportion to their micro-com- studies (e.g., Maxson and Wilson, 1975) plement fixation (MC'F) titers, and all re- but somewhat higher than in one study ported results were obtained with these of salamanders (Maxson et al., 1979). The pooled antisera. data in Table 3 were averaged and a den- Reactivity was measured by the quan- drogram (Fig. 1) was constructed using titative MC'F technique (Champion et al., the method described by Farris (1972), 1974), and the data are reported in im- but modified so that each monophyletic munological distance units (IDU). One group of the tree uses only the closest (in IDU is approximately equivalent to one a cladistic sense) outside lineage in as- amino acid difference (Maxson and Wil- signing relative length to limbs. This son, 1974). modification minimizes errors of infer- ence (Maxson and Wilson, 1975). The RESULTS percent 'standard deviation," defined by Antisera were successfully prepared Fitch and Margoliash (1967) as a measure for all seven species of salamanders used of "goodness of fit" of the tree to the data, in this study. The antibody titers (as de- is 11.4%. The F-statistic defined by Pra- fined in Champion et al., 1974) ranged ger and Wilson (1978) is 6.9%. The meth- from a low of 1500 (P. bellii) to a high of od of tree construction that we have used 6300 (C. multidentatus). The average ti- makes no assumption concerning rates of ter (2750) was somewhat lower than that albumin evolution. typically found in antisera prepared to In addition to our reciprocal tests, uni- albumin of diverse (Maxson directional tests were made using the 5 et al., 1979); this probably is a function antisera of Pseudoeurycea with 13 addi- of the relatively small amounts of albu- tional species of the genus, and the 2

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10 P. brunnato 6 s 8 P sm/f/il 4 P. leprosa 3 PPbe//i P cepha/ica C. bromeliocia C. mu/fidenta/us

L Il l l l II 60 50 40 30 20 10 0 Average ID FIG. 1.-Dendrogram constructed from averaged reciprocal data in Table 3 according to method of Farris (1972), modified as outlined in text. This phylogeny gives the branching order of the lineages leading to the modern species. The numbers on the branches are the amounts of albumin change (expressed in immunological distance units) estimated to have occurred along each branch since the species last shared a common ancestor.

TABLE 4.-Albumin comparisons involving additional species of Pseudoeurycea.

Immunological distance

Species Anti-R Anti-C Anti-L Anti-S Anti-B P. rex species-group P. brunnata (R) 0 72 48 15 53 P. exspectata 27 92 38 18 53 P. goebeli 15 65 36 18 50 P. rex 24 >130 35 20 >105 P. cephulica species-group P. cephalica (C) 49 0 64 62 78 P. altamontana 28 73 54 31 57 P. cochranae 21 74 53 30 68 P. scandens 97 50 P. werleri 61 102 53 65 P. leprosa species-group P. leprosa (L) 53 88 0 35 72 P. juarezi 27 72 31 17 46 P. nigromaculata 53 42 49 P. robertsi 40 85 59 29 59 P. gadovii species-group P. smithi (S) 21 71 35 0 47 P. gadovii 5 79 46 14 58 P. melanomolga 8 4 45 P. unguidentis 8 79 34 0 44 P. bellii species-group P. bellii (B) 39 74 52 43 0

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TABLE 5.-Albumin comparison involving antisera leprosa and cephalica groups. Pseudoeu- to Chiropterotriton. rycea juarezi and P. robertsi of the le- Anti- Anti- prosa group are closer to P. brunnata and Species tested multidentatus bromeliacia P. smithi than they are to P. leprosa, and Chiropterotriton a the shortest distance of any species to P. C. multidentatus 0 98 leprosa is 31 (P. juarezi). The next closest C. chiropterus 53 81 species to P. leprosa (30-40 IDU) are C. chondrostega 29 74 C. dimidiatus 25 members of the gadovii and rex groups: C. priscus 26 71 P. unguidentis, P. smithi, P. rex, P. goe- Chiropterotriton 8 beli, P. exspectata. All are closer to P. le- C. bromeliacia 104 0 prosa than to any member of the leprosa C. cuchumatanus 21 group, save P. juarezi. C. rabbi 90 14 The cephalica group is even more het- C. nasalis 108 64 erogeneous, for none of the members we C. veraepacis 100 67 have tested has an immunological dis- tance less than 70 to P. cephalica, and all antisera of Chiropterotriton with 8 ad- have lesser immunological distances to ditional species of that genus. Results are all other Pseudoeurycea antisera used. reported in Tables 4 and 5. Pseudoeurycea cephalica appears to The species of Pseudoeurycea display have the most differentiated albumin a surprisingly heterogeneous pattern. Im- (range of immunological distances 49- munological distance of P. bellii aver- 130, mean 76) tested in the genus. Two aged 49 + 6 units (SD) to the species of putative members of the cephalica the gadovii group, 52 + 2 units to the group, P. altamontana and P. cochranae, species of the rex group, 59 ? 13 units to have albumin closest to P. brunnata and the leprosa group, and 68 ? 10 units to P. smithi. Our data are incomplete for P. the cephalica group. The bellii group scandens and P. werleri, but both seem contains but a single species; it is essen- to have distinct albumin (minimal dis- tially equidistant from the gadovii and tance of both species to P. leprosa is rex groups, and somewhat further from about 50). the leprosa and cephalica groups. The If P. smithi and P. brunnata are con- members of the gadovii and rex groups sidered part of the same lineage, one are relatively close, but they display sur- might expect relatively unrelated species prising variability considering the rela- to show similar immunological distances tively small immunological distances to both species. For some species this is (12 + 8). Further, P. exspectata and P. indeed the case, and the difference be- rex, of the rex group, are less distant from tween the highest and lowest measure to P. smithi than they are from P. brunnata. P. smithi and P. brunnata is 20% or less Conversely, P. gadovii, of the gadovii of the lowest figure (P. altamontana, av- group, is closer to P. brunnata than to P. erage distance 29.5; P. werleri, 63; P. ni- smithi. The greatest distance shown by gromaculata, 51; P. bellii, 41). The only any member of either of these two groups species that seems markedly less distant is 27 (P. exspectata-P. brunnata), and from one than from the other is P. leprosa three species of the gadovii group (P. ga- (average immunological distance to P. dovii, P. melanomolga, P. unguidentis) smithi, 35; to P. brunnata, 50.5). have very low distances to both P. brun- Immunological distances between the nata and P. smithi (all but one measure- a and ,8 Chiropterotriton are greater than ment less than 10 IDU). those within Pseudoeurycea. The dis- The relatively close similarity of mem- tance between C. multidentatus (a) and bers of the gadovii and rex groups stands C. bromeliacia (,3) is large, with very in sharp contrast to the heterogeneous good reciprocity (98-104).

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Unidirectional tests using the two Chi- smithi group) in the gadovii group, but ropterotriton antisera were made to eight otherwise reached systematic conclu- other species in the genus (Table 5). Im- sions identical to those of Taylor. The munological distance of C. multidenta- only recent systematic account of the en- tus to 8 species ranges from 90-108 (x = tire genus is the provisional annotated 100.5). Immunological distance of C. list of species groups and species by bromeliacia to a species ranges from 71- Wake and Lynch (1976), who observed 90 (x = 82). Within the a group, C. mul- that Pseudoeurycea "is in need of taxo- tidentatus is about equally distant from nomic revision." They essentially fol- C. chondrostega, C. dimidiatus, and C. lowed Taylor, as modified by Baird, ex- priscus (25-29), but the distance from C. cept for assigning Guatemalan species multidentatus to C. chiropterus is con- included by Taylor in the leprosa group siderably greater (53). The ,B group is to a separate rex group. more diverse. The distance of C. bro- The results of the present study sup- meliacia to other /3 species ranges from port the removal of the Guatemalan 14 (C. rabbi) and 21 (C. cuchumatanus) species from the leprosa group, but the to 64 and 67 (C. bromeliacia to C. nasalis immunological data also indicate a close and C. veraepacis, respectively). relationship between the rex and gadovii The mean of reciprocal immunological groups. In view of the lack of morpholog- distance between C. bromeliacia and the ical diagnosis for the rex group, we sug- five antisera of Pseudoeurycea is 77.8 gest that it simply be included within an (range 53-92, SD 14.3). The comparable expanded gadovii group. value for C. multidentatus is similar A "core" group of species has immu- (mean 84.5, range 57-107, SD 14.7). Both nological distances to both P. smithi and C. bromeliacia and C. multidentatus are P. brunnata on the order of 20 or less. more similar to Pseudoeurycea than to These species include P. smithi, P. brun- each other (mean IDU = 101). nata, P. gadovii, P. goebeli, P. melano- molga, and P. unguidentis. Another DISCUSSION group is slightly more distant, with av- Prior to Taylor (1944), most workers erage immunological distances on the or- classified plethodontid salamanders from der of 30: P. exspectata, P. juarezi, P. rex, the New World tropics in the genus Bo- and P. cochranae. Pseudoeurycea robert- litoglossa (= Oedipus). Taylor estab- si and P. altamontana are even more dis- lished both Pseudoeurycea and Chirop- tant (maximum distance = 40). All of the terotriton, which were distinguished aforementioned species are closer to P. mainly on the basis of size (Pseudoeury- smithi and P. brunnata than to any other cea is the larger) and foot shape (Chirop- species for which antisera were tested. terotriton has relatively larger, broader Some of these species have never been feet, with expanded terminal phalanges placed in either the rex or gadovii and a relatively long fifth toe), as well as groups, but all six of the above-listed some osteological characters. To this day "core species have. Pseudoeurycea the genera are recognized essentially as cochranae was placed in the leprosa defined by Taylor, although a number of group by Taylor (1944) and in the ce- additional species have been described. phalica group by Wake and Lynch Taylor (1944) recognized (but did not (1976), whereas our study suggests that diagnose) five species-groups of Pseu- this species is not closely related to doeurycea: gadovii, smithi, cephalica, either lineage. Based on external appear- bellii, and leprosa. A detailed anatomical ance and ecological characteristics, P. study of Pseudoeurycea was presented robertsi and P. altamontana were placed by Baird (1951), who included P. smithi in the leprosa group by Taylor. Wake and (the sole representative of Taylor's Lynch concurred with respect to P. rob-

This content downloaded from 136.152.142.101 on Wed, 03 Feb 2016 22:17:42 UTC All use subject to JSTOR Terms and Conditions June 1981] HERPETOLOGICA 115 ertsi, but assigned P. altamontana to the Wake and Lynch (1976) and Lynch and cephalica group. The present study in- Wake (1978) previously recognized two dicates that both species are far closer to very distinct species-groups in Chirop- P. smithi and P. brunnatathan they are terotriton /8: bromeliacia and picadoi to P. leprosa, and both are remotely re- (Lynch and Wake, 1978, erroneously re- lated (over 70 IDU) to P. cephalica.Pseu- ferred to the latter as the nasalis group). doeuryceajuarezihas been thought to be This arrangement is supported by our a close relative of P. firscheini and P. ni- data. Chiropterotriton bromeliacia, C. gromaculata(Regal, 1966), both of which cuchumatanus and C. rabbi (bromelia- are considered close relatives of P. lepro- cia group) are closely related; the latter sa. Our data indicate that P. juarezi is in- species are 21 and 14 IDU from C. bro- deed reasonably close to P. leprosa (31), meliacia. However, C. nasalis and C. but closer still to P. brunnata(27) and P. veraepacis (picadoi group) are remote smithi (17). from C. bromeliacia (IDU = 64 and 67 Of the species of the leprosagroup that respectively), and presumably from the we tested, only P. nigromaculatais clos- rest of the group as well. The latter two er to P. leprosa than to any other anti- species are only a little closer to C. bro- serum we prepared, but even this rela- meliacia than are some species of Chi- tionship is relatively remote (IDU = 42). ropterotriton a, but relative to C. multi- The content of the cephalica group is dentatus all members of Chiropterotriton brought into question by our discovery /3 are about equally remote. that no member of the cephalica group The dendrograms presented by Wake is closer than 73 IDU to P. cephalica. In- and Lynch (1976) indicate that Chirop- complete data for P. scandens and P. terotriton a is closer to Pseudoeurycea werleri indicate that neither species is than either is to Chiropterotriton /3. Our closer than 50 IDU to any other anti- data indicate that the two antisera of Chi- serum tested. ropterotriton are closer to Pseudoeury- Pseudoeurycea bellii, the only member cea than to each other, but we cannot of the bellii group, is relatively remote with confidence say which is the closer. from all other Pseudoeurycea(smallest The data support the concept of a and / IDU = 43, to P. smithi). groups, and also support the contention We conclude that most of the species of Lynch and Wake (1978) that the bro- of Pseudoeuryceatested are moderately meliacia and picadoi groups are very dis- close relatives that provisionally are to be tinct. The implications of these results considered members of the gadovii will be discussed elsewhere, in the species-group. Our data indicate that the con- text of a detailed morphological bellii, leprosa, and cephalica groups analysis. each contain a single, phyletically dis- Rabb (1956) described C. priscus, a tinct species. species he believed to be somewhat in- Wake and Lynch (1976) recognized termediate in anatomical features be- three species-groups within Chiroptero- tween Chiropterotriton and Pseudoeu- triton a: chiropterus, multidentatus, and rycea. He specifically compared C. priscus. Our immunological data indi- priscus with P. galeanae, a member of cate that two species-groups may be jus- the cephalica group. Chiropterotriton tified. Chiropterotriton chondrostega, C. priscus is immunologically close to C. dimidiatus (both previously assigned to multidentatus (IDU = 26), but remote the chiropterus group), and C. priscus from P. cephalica (IDU = 74). Rabb are all relatively close to C. multidenta- (1956) and Wake and Lynch (1976) hy- tus, whereas C. chiropteruscomprises a pothesized that the similarities between second group that is relatively remote C. priscus and some Pseudoeurycea are from C. multidentatus. based on retained generalized and prim-

This content downloaded from 136.152.142.101 on Wed, 03 Feb 2016 22:17:42 UTC All use subject to JSTOR Terms and Conditions 116 HERPETOLOGICA [Vol. 37, No. 2 itive anatomical features. Our data are FITCH, W. M., AND E. MARGOLIASH. 1967. Con- consistent with such an interpretation. struction of phylogenetic trees. Science 155:279- As is evident from inspection of the 284. HARTMAN, B. K., AND S. UDENFRIEND. 1969. A internodal distances in Figure 1, rates of method for immediate visualization of proteins in albumin evolution in this assemblage acrylamide gels and its use for preparation of an- have been reasonably regular. An excep- tibodies to enzymes. Anal. Biochem. 30:391-394. tion is the lineage containing P. brun- LARSON, A., D. B. WAKE, L. R. MAXSON, AND R. HIGHTON. 1981. A molecular phylogenetic per- nata and P. smithi, which has accumu- spective on the origins of morphological novelties lated considerably less immunological in the salamanders of the tribe Plethodontini distance during its evolution than have (Amphibia, Plethodontidae). Evolution, in press. the other lineages. While the accumula- LYNCH, J. F., AND D. B. WAKE. 1978. A new tion of immunological distance is species of Chiropterotriton (Amphibia: Caudata) mainly from Baja Verapaz, Guatemala, with comments on a time-dependent phenomenon (Wilson relationships among Central American members et al., 1977), at least some species of of the genus. Contrib. Sci. Nat. Hist. Mus. Los Pseudoeurycea have undergone different Angeles Co. No. 294. rates of change. If we accept the estimate MAXSON, L. R., R. HIGHTON, AND D. B. WAKE. 1979. Albumin evolution and its phylogenetic that roughly 100 units of immunological implications in the plethodontid salamander gen- distance accumulate between two lin- era Plethodon and Ensatina. Copeia 1979:502- eages every 55-60 million years (Maxson 508. and Wilson, 1975; Wilson et al., 1977), MAXSON, L. R., AND R. D. MAXSON. 1979. Com- the evolutionary of some of parative albumin and biochemical evolution in history the plethodontid salamanders. Evolution 33:1057- groups discussed herein may extend over 1062. most of the Cenozoic. The species of MAXSON, L. R., AND A. C. WILSON. 1974. Conver- Pseudoeurycea and Plethodon show very gent morphological evolution detected by study- similar distributions of immunological ing proteins of tree frogs in the Hyla eximia species group. Science 185:66-68. distances (Maxson and Maxson, 1979; . 1975. Albumin evolution and organismal Maxson et al., 1979), indicating that the evolution in tree frogs (Hylidae). Syst. Zool. 24: 1- two genera, neither of which is strongly 15. differentiated in morphology, may be of PRAGER, E. M., AND A. C. WILSON. 1978. Con- similar age and have had a long history struction of phylogenetic trees for proteins and nucleic acids: empirical evaluation of alternative of cladogenesis. matrix methods. J. Mol. Evol. 11:129-142. RABB, G. B. 1956. A new plethodontid salamander Acknowledgments.-We thank James Lynch for from Nuevo Leon, Mexico. Fieldiana Zool. encouragement, assistance with field work, and 39: 11-20. careful evaluation of the manuscript. Marvalee REGAL, P. J. 1966. A new plethodontid salamander Wake and Stanley Sessions offered helpful com- from Oaxaca, Mexico. Am. Mus. Novit. No. 2266. ments. Field assistance was also provided by Ste- TAYLOR, E. H. 1944. The genera of plethodont sal- phen Busack, Paul Elias, James Hanken, Erik Ko- amanders in Mexico, pt. 1. Univ. Kans. Sci. ford, Theodore Papenfuss, Bull. Stanley Sessions, and 30:189-232. Thomas Wake. Technical assistance was provided WAKE, D. B. 1966. Comparative osteology and evo- by Ellendee Pepper and Gloria Wurst. The research lution of the lungless salamanders, family Pleth- was supported by NSF grants DEB 78-03008 (to odontidae. Mem. South. Calif. Acad. Sci. 4: Wake) and DEB 78-23396 (to Maxson). 1-111. WAKE, D. B., AND J. F. LYNCH. 1976. The distri- bution, LITERATURE CITED ecology, and evolutionary history of plethodontid salamanders in tropical America. BAIRD, I. L. 1951. An anatomical study of certain Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. No. salamanders of the genus Pseudoeurycea. Univ. 25. Kans. Sci. Bull. 34:221-265. WAKE, D. B., L. R. MAXSON, AND G. Z. WURST. CHAMPION, A. B., E. M. PRAGER, D. WACHTER, AND 1978. Genetic differentiation, albumin evolution, A. C. WILSON. 1974. Micro-complement fixation. and their biogeographic implications in pletho- Pp. 397-416. In C. A. Wright (Ed.), Biochemical dontid salamanders (genus Hydromantes) of Cal- and Immunological of . Aca- ifornia and southern Europe. Evolution 32:529- demic Press, New York and London. 539. FARRIS, J. S. 1972. Estimating phylogenetic trees WALLACE, D. G., M. C. KING, AND A. C. WILSON. from distance matrices. Am. Nat. 106:645-688. 1973. Albumin differences among ranid frogs:

This content downloaded from 136.152.142.101 on Wed, 03 Feb 2016 22:17:42 UTC All use subject to JSTOR Terms and Conditions June 1981] HERPETOLOGICA 117

taxonomic and phylogenetic implications. Syst. Department of Genetics and Develop- Zool. 22:1-4. ment, University of Illinois, Urbana, IL WILSON, A. C., S. S. CARLSON, AND T. J. WHITE. 61801, USA, and Museum of Vertebrate 1977. Biochemical evolution. Ann. Rev. Bio- chem. 46:573-639. Zoology, University of California, Berke- ley, CA 94720, USA Accepted: 8 December 1980

Herpetologica, 37(2), 1981, 117-121 ? 1981 by The Herpetologists' League, Inc.

IN MEMORIAM: ARTHUR LOVERIDGE

CARL GANS

ON 16 February 1980 the herpetologi- had over 300 cases of natural history and cal community lost Arthur Loveridge- anthropological specimens and over 250 professional herpetologist, explorer, pop- jars of "spirit-preserved" reptiles. He ular writer, radio personality, soldier, and noted proudly that he was prepared to long-time curator of amphibians and rep- handle techniques of mounting and pre- tiles at the Museum of Comparative Zo- serving all of his captures, described his ology. He published his first paper in complex system of registration and col- 1913 and was still actively concerned lecting numbers, and referred to an 80- with natural history at the time of his page catalog, with short descriptions of death on the island of St. Helena. In be- all specimens, all typed by himself. Even tween, he published well over 200 books, his handwriting was small and meticu- research papers, popular articles, and re- lously formed with little change between views, including the volumes Many Hap- top and bottom of pages or successive py Days I've Squandered (1944), Tomor- sheets. Field notes and labels (as well as row's a Holiday (1944), Reptiles of the his many letters) alike were beautifully Pacific World (1945), I Drank the Zam- readable. besi (1953), and Forest Safari (1956). Judging from comments in his books Mr. Loveridge was born in Penarth, and from documents made available from Glamorgan, Wales, on 28 May 1891. He the archives of Harvard University, Love- appears to have had an abiding interest ridge completed school and then was in natural history from his earliest years. made by his family to serve two years in An enthusiasm for natural history is not a business apprenticeship, where he was uncommon in boys, nor is the trend to in charge of the storeroom and associated assemble masses of natural history ob- record keeping. He next obtained a one- jects. The spectacular difference be- year course at the University College of tween Loveridge and other enthusiastic South Wales, Cardiff, under Professor W. boy-naturalists was his total commitment N. Parker, and then took a six-month ap- to the pursuit of natural history as a ca- pointment as an assistant in the Man- reer. In 1914, at the age of 24, he applied chester University Museum, cataloging, for the curatorship of the Nairobi Mu- rebottling, and re-identifying the collec- seum, advising in the application that he tion. In 1911, he was appointed assistant

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