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Analisis filogenetico en el grupo Pristimantis Unistrigatus (Anura, craugastoridae) basado en caracteres morfologicos.

PHYLOGENETIC ANALYSIS WHITHIN THE PRISTIMANTIS UNISTRIGATUS (ANURA, CRAUGASTORIDAE) GROUP BASED ON MORPHOLOGICAL CHARACTERS

INTRODUCTION

Frost et al. (2006) doubted the existence of the genus Eleutherodactylus Dumeril and Bibron, 1841 as monophyletic, and suggested that it should be divided into many monophyletic genera. Heinicke et al. (2008) divided the former genus Eleutherodactylus plus the genus Brachycephalus Fitzinger, 1826 in three Clades: the South American Clade, the Caribbean Clade and the Middle American Clade. The Eleutherodactylus unistrigatus group was included in the South American Clade using the available name Pristimantis Jimenez de la Espada, 1870. This group is one of the richest Pristimantis groups, with members occurring in the Andes and the Amazonian basin, ranging from Bolivia to Colombia (Hedges et al. 2008, Padial et al. 2009).

In the literature related to the diagnosis of the unistrigatus group species of the Cordillera Oriental of Colombia, potential kinship and characteristics in common are referred (Duellman & Simmons 1977, Lynch 1973, 1978, 1981, 1983, 1984, 1994a, 1994b, 1998, 2003, Lynch & Duellman 1980, Lynch & Duellman 1997, Lynch & Suarez 2002, Pyburn & Lynch 1981). However, there have been no formal relationship hypotheses supported by cladograms.

It is perhaps for this Pristimantis group that were proposed the highest number of hypotheses of relationships in Colombia, because it is the richest in species number in this country. From the perspective of phylogenetic systematics of the genus Pristimantis, the work that has a growing number of Colombian species for the unistrigatus group (sensu Lynch & Duellman 1997) of the Cordillera Oriental of Colombia has been that of Galindo (2004), based on external characters. However, the papers of Duellman & Simmons (1977), Lynch (1973, 1975a, 1975b, 1978, 1981, 1983, 1984, 1994a, 1994b, 1998, 2003), Lynch & Duellman (1980), Lynch & Suarez (2002) and Pyburn & Lynch (1981) have revealed potential relationships for species of the unistrigatus group, some with formal cladograms, but the majority without them. Lynch (1984) proposed a hypothesis of relationship for the unsitrigatus group without including any of the species used here, but some belonging to the "assemblages" pyrrhomerus and myersi, suggesting these should be combined by a series of synapomorphies found in external characters. Finally, recently Padial et al. (2009) considered that the Pristimantis unistrigatus group is not monophyletic.

Other groups, which have launched hypotheses of relationship, either with or without explicit cladograms are: discoidalis (Lynch 1989) (with explicit cladogram), using external and cranial characters, and sulcatus (Lynch 1997) (with explicit cladogram), finding for these relationships nine characters from the skull. As this paper aims to use morphology as a fundamental element to establish kinship hypothesis, the discussion will not be based on specific morphological differences between what we found and what other authors have found, but between the relationships proposed here based on potential synapomorphies, and other hypotheses and characters used to construct them.

From morphology's point of view, the contributions are all innovative in the sense that no previous research has been dedicated solely to the study of muscles of hands and feet for the phylogenetic analysis of the Pristimantis genus. Similarly, although there are some previous descriptions about hand muscles of some species of the genus (e.g.Burton 1998a, Dunlap 1960), these were mainly for comparative purposes (Dunlap 1960, Salgar et al. 2009), or with the aim to find characters to establish hypotheses of relationships (Burton 1998a) but there was not a genus-specific analysis performed.

Previous studies in the literature, using both muscles of hands and forearms and feet to propose phylogenetic relationships in frogs are very rare; in fact, there is only one (Faivovich 2002), but in the genus Scinax (Hylidae). The most comprehensive descriptive work in this regard have been those of Gaup (1886), Dunlap (1960), Andersen (1978) and, more recently, Burton (1996, 1998a,b, 2001, 2004). Most of them propose (tacitly or explicitly) some assumptions about the potential systematic value of characters taken from hands and feet in anuran amphibians, without reaching cladistic analysis themselves. Only in one of his most recent work, Burton (2004) developed a cladistic analysis using only the muscles of the feet in the family Hylidae. The only existing descriptions of this musculature in Colombian species of frogs has been conducted by Salgar (2003) and Salgar et al. (2009). Although these last two analyses are brief, for the first time, possible synapomorphies for the genus Pristimantis (intrageneric relationships) based on the muscles of hands and feet were revealed. Lynch (1984) proposed some characteres but for relationships between species groups, based on osteological characters and some of the muscles (but not hands and feet).

Hedges et al. (2008) redefined the Pristimantis unistrigatus group changing the species included to it, assigning 193 species to this group, changing some of the species previously identified into the group by Lynch (1976a, Lynch & Duellman 1997, Duellman & Pramuk 1999), and by Lynch & Duellman (1997) and Acosta (2000) for Colombia.

Taking into account that there are two different classifications for the Pristimantis unistrigatus group, our aim is to compare our results in using morphological characters from hand and foot muscles in both proposals, because there are not enough consensus about which is the best classification. It is for this reason that both the number of species and characters in both analyses differs (see results). The only classification that include all species of our study is that by Lynch & Duellman (1997). The number of species in classification by Hedges et al. (2008) is lower, because they do not consider some of the species by Lynch & Duellman (1997) as belonging to the P unistrigatus group. Owing to this account, the number of characters also changed. In comparing our results with those by Heinicke et al. (2008), and Hedges et al. (2008), we can conclude that they agree that P. unistrigatus group is not monophyletic.

Materials and methods

A total of 42 adult specimens (Table 1) representing 17 species of Pristimantis of the Cordillera Oriental of Colombia and one of Craugastor were dissected. Specimens belong to the Museo de Historia Natural (MUJ) of the Pontificia Universidad Javeriana in Bogota, Colombia, and to the Instituto de Ciencias Naturales (ICN) of the Universidad Nacional de Colombia in Bogota. Taking into account the species belonging to the unistrigatus group sensu Hedges et al. (2008), we decided to compare results considering the former classification made by Lynch & Duellman (1997).We chose the following two ingroups: sensu Lynch & Duellman (1997) (LD): Pristimantis anolirex (Lynch, 1983): ICN 33522, 15444; P bacchus (Lynch, 1984): ICN 33164, 33170, 34232; P bicolor (Rueda-Almonacid and Lynch, 1983): ICN 14474, 26321; P bogotensis (Peters, 1863): MUJ 052, 465, 481, 687; P. elegans (Peters, 1863): MUJ 036; P frater (Werner, 1899): ICN 40579, 49911; P. jorgevelosai (Lynch, 1994): ICN 15299, 15318; P lynchi (Duellman and Simmons, 1977): MUJ 1480, 1538, 1507, 2036; ICN 01932, 01937; P merostictus (Lynch, 1984): ICN 34233, 34235; P miyatai (Lynch, 1984): ICN 50062, 50065, 50069; P nervicus (Lynch, 1994): MUJ 023, 1176; P nicefori (Cochran and Goin, 1970): ICN 20994, 21119; P prolixodiscus (Lynch, 1978): ICN 10104, 10107; P pugnax (Lynch, 1973): ICN 22981, 23187; P spilogaster (Lynch, 1984): ICN 12445, 12446; P. taeniatus (Boulenger, 1912): MUJ 2452, 2420; P. tamsitti (Cochran and Goin, 1970): ICN 22949, 22951, 23638; Sensu Hedges et al. (2008) (HEA): P anolirex (Lynch, 1983), P bacchus (Lynch, 1984), P bogotensis (Peters, 1863), P elegans (Peters, 1863), P lynchi (Duellman and Simmons, 1977), P. merostictus (Lynch, 1984), P. nervicus (Lynch, 1994), P. nicefori (Cochran and Goin, 1970), P pugnax (Lynch, 1973), P. spilogaster (Lynch, 1984), P. taeniatus (Boulenger, 1912), P. tamsitti (Cochran and Goin, 1970).

Outgroups: Craugastor raniformis: MUJ 2871, 2689 for the LD matrix, and P frater for the HEA matrix (Tables 2 and 3). The new classification proposed by Hedges et al. (2008) for some of the species formerly assigned to unistigatus group is as follows: P. frater: frater group; P bicolor: species series P. (Hypodiction) ridens; P.miyatai: P. frater group; P. jorgevelosai: species series P. (Hypodiction) ridens; and P. prolixodiscus: lacrimosus group.

Muscles

We observed about 90 small hand and feet muscles, both superficial and deep, of which about 50 superficial were chosen. For choosing the characters and states, we did comparisons between muscles from ten species of Pristimantis used for the study with respect to different aspects that are related below. Dissections were done in hands, feet, and legs and forearms in order to examine those muscles associated morphologically with the hands and feet. We used a dissecting microscope and a reversible Weigert's iodine solution that stains muscles (Bock & Shear 1972) to see the direction of the fibers, obtaining a clear differentiation of muscles and tendons, including the origins and insertions. For all specimens both right and left hands and feet superficial muscles were dissected and examined, following the protocols proposed by Raikow et al. (1990). For each muscle we identified: a. origin; b. insertion; c. spatial arrangement with respect to the hand and foot, and d. spatial relationship with adjacent muscles. Although we explored mainly superficial musculature, we also dissected some deep muscles considering that we took into account characters proposed by Burton (1998a, 2004). The nomenclature was taken from that proposed by Dunlap (1960), Andersen (1978), Burton (1998a, 2004), and Salgar et al. (2009). We used for most cases, at least one male and one female of each species to the extent that the number of specimens of the collection allowed it, to determine the existence of sexual dimorphism. Finally, we drew pictures of characters considered as potential synapomorphies using photographs and the Adobe Illustrator software. For the accuracy of the origin and insertion of muscles, we made use of clearing and staining skeletons of P. bogotensis.

External characters

These were taken from taxonomic descriptions after Cochran & Goin (1970), Lynch (1983), Lynch (1984), Rueda & Lynch (1983), Dunn (1944), Hoyos (1991), Pyburn & Lynch (1991), Lynch (1994), Duellman & Simmons (1997), Lynch (1998), Lynch (1978), and Lynch & Duellman (1980), and after our own observations.

Phylogenetic analysis

For the phylogenetic analysis, we compared each result taking into account two matrices (see below). We did heuristic searches in TNT, performing TBR branch swapping on 100 addition replicates. Matrices were running considering characters with the same weight. We did not find many intraspecific (polymorphic) variations or many multistate characters, however we use all of those we found for the phylogenetic analyses. Polymorphic characters were coded according to Faivovich (2002) as follows: A = 0/1; B = 0/2, and C = 1/2, and multistate characters were all considered nonadditive and unordered. We did not get whether retention (RI) or consistency indices (CI), given that we are comparing groups of different sizes (e.g. Conrad 2008), so the value for the CI will be greater in the HEA (Sensu Hedges et al., 2008) group than in the LD (sensu Lynch & Duellman, 1997) group because we considered less species in the first group than in the second one, and the RI will be less in the HEA for the same reason.

We choose one outgroup species for each analysis, following recommendation by Nixon & Carpenter (1993: 419) about the fact that "...a third pervasive myth, the notion that more than one outgroup is somehow required for correct polarity determination", and taking into account that "Outgroups and polarity are the same problem" (Nixon Nixon & Carpenter 1993: 420). We only show the unambiguous synapomorphies.

Results

We scored 81 characters, from which 64 are muscle characters (44 from the manus and 20 from the pes), and 17 external characters. Four manus characters were taken from Burton (1998a), and five pes characters from Burton (2004). Based on this matrix, we eliminated those characters according to changes in the taxa for each matrix as we describe below.

Phylogenetic analysis

LD P. unistrigatus group

By eliminating those characters with only one character state present in the outgroup and ingroup, we took 70 from the 81 characters (Appendix 1). The cladistic analysis resulted in one fully resolved most-parismonious tree (Fig. 1) with a length of 202. We found character states 62(0) (First finger shorter than second) and 23(1) (The origin of the m. adductor policis is not covered by the aponeurosis palmaris) (Fig. 2) as the only two unambiguous putative synapomorphies. The first one included almost all species leaving away P nicefori, and the second one is a character shared by P prolixodiscus and P bicolor.

HEA P. unistrigatus group (Appendix 2)

We took 60 characters for the analysis (Appendix 2). The cladistic analysis resulted in 20 most-parismonious trees of 139 steps. The strict consensus tree (Fig. 3) shows a node 14 not resolved, consisting of taxa P lynchi, P nervicus, P elegans, P bogotensis, P nicefori, P. anolirex, P. bacchus, and P. taeniatus. The only unambiguous putative synapomorphy (found in all trees) is the character state 44(1) (Origin of the tendo superficialis hallucis from the aponeurosis plantaris including some fibers of the m. lumbricalis brevis hallucis) (Fig. 4) relating P. spilogaster and P. merostictus.

DISCUSSION

As shown in the cladograms obtained for both unistrigatus groups, the LD (Fig. 1) group is more resolved than the strict consensus of the HEA (Fig. 3), but there are no synapomorphies that support it, except partially for nodes 21 and 32 in LD, and exclusively for node16 in HEA, this means that all topologies are weakly supported by synapomorphies. In node 22 of DL no synapomorphies were found, whereas for the other nodes, potential synapomorphies are actually reversals or convergences.

Taking into account the results and species considered by HEA and LD, we could say that the assigned HEA unistrigatus group is weaker because the relationships are less resolved, even if the number of steps is smaller (it has a smaller number of taxa). On the other hand, even with a larger number of species involved, the proposed LD unistrigatus group can be considered stronger if we take the fact that we have got a single tree completely resolved; this does not mean that we are clear about the putative synapomorphies involved, as we saw above. It is remarkable that the relationships between P. pugnax and P. tamsitti, and between P. spilogaster and P merostictus are retained in both cladograms, meaning that these species maybe considered sisters, but without finding putative synapomorphies to support the relationship between P pugnax and P. tamsitti.

Limb muscles and phylogeny

All characters and character states obtained were chosen bearing in mind character distribution and conditions deemed to be worth for the analysis. Burton's (1998a, 2004) characters and character states were used just as he defined them, but in some cases we believed that some had to be redefined due to the fact that they were not clearly named. Then, we think that most of these characters are new and very useful for future analyses in frog morphology and phylogeny studies. It is fascinating how many characters have found on the hands and feet, although only the superficial muscles have been observed. More characters were found in the hands (44, including four of Burton) than in the feet (21, including six of Burton). Similarly, after making a clearance based on external descriptions and original observations, we obtained 26 characters. Faivovich (2002) used 14 characters from the manus and seven from the pes, from which he found two synapomorphies for Scinax Wagler, 1830, both from the manus. However, although we did not find any useful characters taken from hand and foot musculature, this does not mean that these characters should be dismissed from phylogenetic studies in frogs. Burton (1998a) found some hand muscle character conditions useful to support intrafamiliar relationships in Leptodactylids. Likewise, Burton (1998b) found a hand muscle character condition within Ranidae Rafinesque, 1814, stressing that superficial muscles can vary more than deep ones, but that these muscle can contribute to get frog relationships. It is apparently historically clear that superficial hand muscles may play a role more important to resolve frog relationships than superficial feet muscles, although we found muscles of both that could be putative synapomorphies.

External characters

Phylogenetic analysis including external characters is rare in frogs, although identification of most of the species groups and all species have been based on these characters. The results show that characteristics of the fingers are still important, and can be used in establishing kinship, but were obviously not essential to support the monophyly of the unistrigatus group. Apparently, the other external characters are not useful because, in any case, it was established that they could participate as potential synapomorphies. Maybe we should pay attention to the condition of tympanic annulus and to the dorsal view of the snout because for the two analyses that show that these characters appear as potential synapomorphies (nodes 21 and 23 respectively in LD group, and node 15 in HEA group), although actually they are homoplasies.

To compare our results with those by Heinicke et al. (2008) and Hedges et al. (2008) is difficult, because most of the species included in our work were not used by these authors. However, our results agree in identifying the unistrigatus group as non monophyletic. Heinicke et al. (2008) only included six spececies belonging to the former P unistrigatus group (Lynch, 1976; Lynch et al., 1997). Taking into account this classification, their results showed that this group is monophyletic. However, based on Hedges et al. (2008) classification P. ockendeni, in the P unistrigatus group by Heinicke et al. (2008), belongs to the P frater group; in this sense, the group would become not monophyletic.

CONCLUSIONS

Although the muscles of hands and feet and external characters useful in taxonomy failed to identify the Pristimantis unistrigatus as monophyletic, this does not mean they are not useful for establishing relationships within the genus. It is possible that including more unistrigatus group species in the study, the situation changes, however, it seems also essential to include the deep muscles of the hands and feet and external characters as well. It is possible that other characters facilitate to define relationships among species within the genus. However, it is necessary to explore deeper the relationship P spilogaster-P. merostictus, and P. pugnax-P tamsitti, because we need whether to confirm or falsify if these clades actually are monophyletic. By detecting the difficulties that several researchers (e,g, Hedges et al. 2008; Padial et al. 2009) and our own study have found to discover the monophyly of the P unistrigatus group and other ones in frogs, the question we pose is: Is it worthwhile to keep this informal taxonomic hierarchy called group in systematics of frogs?

Appendix 1. List of characters and character states for the LD matrix.

Manus

1. Number of insertions of the m. extensor digitorum communis longus

0 Four

1 Three

2 Two

2. Number of origins of the m. epicondylocubitalis

0 Two, one slip in common with the m. extensor carpi ulnaris and the other slip free.

1 One slip, not in common with the m. extensor carpi ulnaris

3. Relationship between the m. epicondylocubitalis and the m. epitrochleocubitalis

0 Bipennated

1 Parallel

4. Type of origin of the m. epitrochleocubitalis

0 Double

1 Single

5. Insertion of the m. epitrochleocubitalis

0 Not covered by the m. palmaris brevis

1 Covered by the m. palmaris brevis

6. Insertion of the m. epicondylocubitalis in relation with the m. palmaris brevis

0 It does not cover the m. palmaris brevis

1 It covers the m. palmaris brevis

7. Relationship between the m. epitrochleocubitalis and the external slip of the m. palmaris longus

0 The m. palmaris longus is partially covered by the m. palmaris longus

1 The m. palmaris longus is not covered by the m. palmaris longus

8. Relationship between the m. epitrochleocubitalis and the m. epicondylocubitalis

0 The fibers fuse partially on the external surface of the radioulna

1 The fibers completely fuse on the external surface of the radioulna

9. Relationship between the internal slip of the m. extensor digitorum communis longus and the m. extensor carpi radialis

0 Proximally partially fused

1 Completely fused

2 Not fused

10. Relationship betweem the m. extensor brevis superficialis Digiti V and the m. abductor brevis Digiti V

0 They are partially fused

1 They are not fused

11. Number of insertion of the m. extensor brevis superficialis Digiti V

0 One

1 Two

12. Origin of the m. abductor brevis Digiti V 0 From the distal carpal 5-4-3

1 From the ulnare

13. Presence of the m. abductor ulnocarpalis

0 Present

1 Absent

14. Origin of the m. palmaris longus

0 By two slips

1 By one slip

15. Insertion of the m. palmaris longus

0 Simple

1 Double

16. Connection between the tendons of the m. palmaris longus and the flexor tendons

0 Single tendon that divides to become tendones superficiales (TS) IV and V

1 Two tendons that cross over before becoming TS IV and V

17. Relationship between the fibers of the m. palmaris brevis and the m. palmaris longus

0 The fibers of the m. palmaris brevis are dorsal with respect to the tendons of insertion of the m. palmaris longus

1 The m. palmaris brevis is between the two tendons of insertion of the m. palmaris longus

18. Relationship between the insertion of the m. palmaris brevis and the tendons of insertion of the

m. palmaris longus

0 m. palmaris brevis articulates with the tendons of insertion of the m. palmaris longus

1 The m. palmaris brevis does not articulate with the m. palmaris longus

19. Relationship between the m. palmaris longus and the m.flexor carpi ulnaris

0 They are partially fused

1 They are not fused

20. Origin of the m. adductor pollicis

0 From the ventral surface of the carpal 5-4-3

1 From the internal surface of the distal carpal 5-4-3

21. Insertion of the m. adductor pollicis on the prepollex

0 Adjacent to the insertion of the m. abductor pollicis

1 Covers the insertion of the m. abductor pollicis

22. Relationship between the m. adductor pollicis and the m. abductor pollicis

0 Both muscles are independent

1 They are fused

23. Relationship between the origin of the m. adductor pollicis and the aponeurosis palmaris

0 The origin is covered by the aponeurosis palmaris

1 The origin is not covered by the aponeurosis palmaris

24. Condition of the proximal end of the m. lumbricalis brevis indicis

0 Dorsal to the m. adductor pollicis

1 Contiguous to the m. adductor pollicis

25. Nature of the origin of the tendon of the m. lumbricalis brevis Digiti III

0 Short and wide with respect to the muscle

1 Long and thin with respect to the muscle

26. Origin of the m. lumbricalis brevis Digiti III in relation with the tendo superficialis & caput. profundum Digiti III

0 Dorsal

1 Laterodorsal

27. Presence of fibers of the tendo superficialis & caput. profundum Digiti III

0 Only on the external surface of the tendon

1 On the external and internal surface of the tendon

2 Only on the internal surface of the tendon

28. Nature of the m. lumbricalis brevis Digiti IV

0 Two slips

1 Only one slip

29. Position of the insertion tendons of the m. lumbricalis longus Digiti IV with respect to the tendo superficialis aponeurosis palmaris prodigiti IV

0 Lateral to the tendo superficialis aponeurosis palmaris prodigiti IV

1 The insertion is via the tendo superficialis aponeurosis palmaris prodigiti IV

30. Origin of the m. lumbricalis longus Digiti V

0 From the aponeurosis palmaris and the TS V

1 Only from the TS V

31. Number of insertions of m. lumbricalis longus Digiti V

0 One

1 Two

32. Number of slips that compose the m. lumbricalis brevis Digiti V

0 One

1 Two

Pes

33. Type of insertion of the m. extensor digitorum communis longus

0 Single

1 Multiple

34. Number of insertions of the m. extensor brevis superficialis hallucis

0 Two

1 Three

2 Single

35. Number of insertions of the m. extensor brevis superficialis Digiti II

0 One

1 Two

36. Relationship between the insertions of the m. extensor brevis superficialis Digiti II and the m. extensor digitorum communis longus

0 Common insertion between m. extensor brevis superficialis Digiti II and the internal slip of m. extensor digitorum communis longus

1 Inependent insertions

37. Presence of the m. extensor brevis medius hallucis

0 Absent

1 Present

38. Presence of the m. extensor brevis medius Digiti II

0 Absent

1 Present

39. Number of insertion slips of the m. extensor brevis superficialis Digiti III

0 One

1 Two

40. Relationship between the insertions of the m. extensor brevis superficialis Digiti III and the m. extensor digitorum communis longus

0 Common insertion between m. extensor brevis superficialis Digiti III and any slip of m. extensor digitorum communis longus

1 Independent insertions

41. Origin of the m. extensor brevis superficialis Digiti IV

0 By two independent slips

1 By two proximally fused slips

2 By one slip

42. Presence of the m. extensor brevis superficialis Digiti V

0 Absent

1 Present

43. Origin of the m. abductor prehallucis

0 From the internal surface of the aponeurosis plantaris

1 From the distal internal surface of the tarsus

44. Presence of the m. abductor brevis plantaris hallucis

0 Absent

1 Present

45. Origin of the m. lumbricalis brevis Digiti II

0 From the aponeurosis plantaris

1 From the proximal end of the tendo superficialis hallucis

46. Type of origin of the m. lumbricalis brevis Digiti III

0 Tendinous

1 Fleshy

47. Relationship between the external slip of the m. lumbricalis brevis Digiti IV and the internal slip of m. lumbricalis brevis Digiti V

0 Fused only in the origin

1 Fused beyond its origin

Burton characters

Manus

48. Presence of the internal slip of the m. lumbricalis brevis Digiti III

0 Absent

1 Present

49. Number of slips of the m. lumbricalis longus Digiti IV

0 Two

1 Only one, the internal slip

50. Origin of the m. extensor brevis superficialis Digiti V

0 From both, the ulnare and distal carpals 5-4-3

1 From the ulnare

Pes

51. Origin of the tendo superficialis hallucis

0 From the aponeurosis plantaris

1 From the aponeurosis plantaris including some fibers of the m. lumbricalis brevis hallucis

2 From a muscle that originates from the distal tarsal 2-3

52. Origin of the tendo superficialis Digiti II

0 From the aponeurosis plantaris

1 It serves as the insertion tendon of the m. transversus plantae distalis

53. Origin of the tendo superficialis Digiti III

0 From the m.flexor digiti brevis superficialis only

1 From the aponeurosis plantaris

2 From a superficial tendon that emerges centrally on the plantar surface of the aponeurosis plantaris

54. Number of insertion tendons of the m. lumbricalis logissimus Digiti IV

0 Two

1 One

External characters

55. Skin on venter

0 Areolate

1 Smooth

56. Presence of dorsolateral folds

0 Present

1 Absent

57. Nature of tympannic anulus

0 Complete

1 Incomplete

58. Dorsal view of snout

0 Rounded

1 Subacuminate

2 Acuminate

59. Shape of snout in profile

0 Rounded

1 Truncated

2 Acute

60. Presence of cranial crests

0 Present

1 Absent

61. Width of the upper eyelid in relation with the interorbital distance

0 Narrower than the inter-orbital distance

1 As wide as the inter-orbital distance

62. Length of fingers I and II

0 First finger shorter than second

1 First and second finger equally or almost equally long

2 First finger longer than second

63. Presence of finger pads or discs

0 Present

1 Absent

64. Presence of skin variations on the ulnar region

0 Present

1 Absent

65. Presence of skin variations on the tarsals

0 Present

1 Absent

66. Presence of skin variations on the heel

0 Present

1 Absent

67. Presence of supernumerary plantar tubercles 0 Present

1 Absent

68. Presence of webbed toes

0 Present

1 Absent

69. Presence of toe pads or discs

0 Present

1 Absent

70. Presence of lateral fringes on toes

0 Present

1 Absent

Appendix 2. List of characters and character states for the HEA matrix.

Manus

1. Number of insertions of the m. extensor digitorum communis longus

0 Three

1 Two

2. Relationship between the m. epicondylocubitalis and the m. epitrochleocubitalis

0 Bipennated

1 Parallel

3. Type of origin of the m. epitrochleocubitalis

0 Double

1 Single

4. Insertion of the m. epitrochleocubitalis

0 Not covered by the m. palmaris brevis

1 Covered by the m. palmaris brevis

5. Insertion of the m. epicondylocubitalis in relation with the m. palmaris brevis

0 It does not cover the m. palmaris brevis

1 It covers the m. palmaris brevis.

6. Relationship between the m. epitrochleocubitalis and the external slip of the m. palmaris longus

0 The m. epitrochleocubitalis is partially covered by the m. palmaris longus

1 The m. epitrochleocubitalis is not covered by the m. palmaris longus.

7. Relationship between the m. epitrochleocubitalis and the m. apicondylocubitalis

0 The fibers fuse partially on the external surface of the radioulna

1 The fibers completely fuse on the external surface of the radioulna

8. Relationship between the internl slip of the m. extensor digitorum communis longus. and the m. extensor carpi radialis

0 Proximally partially fused

1 Completely fused

2 Not fused

9. Relationship between the m. extensor brevis superficialis Digiti V and the m. abductor brevis Digiti V

0 They are partially fused

1 They are not fused

10. Number of insertion of the m. extensor brevis superficialis Digiti V

0 One

1 Two

11. Origin of the m. abductor brevis Digiti V

0 From the distal carpal 5-4-3

1 From the ulnare

12. Presence of the m. abductor ulnocarpalis

0 Present

1 Absent

13. Origin of the m. palmaris longus

0 By two slips

1 By one slip

14. Insertion of the m. palmaris longus

0 Simple

1 Double

15. Connection between the tendons of the m. palmaris longus and the flexor tendons

0 Single tendon that divides to become tendones superficiales(TS) IV and V

1 Two tendons that cross over before becoming TS IV and V

16. Relationship between the fibers of the m. palmaris brevis and the m. palmaris longus

0 The fibers of the m. palmaris brevis are dorsal with respect to the tendons of insertion of the m. palmaris longus

1 The m. palmaris brevis is between the two tendons of insertion of the m. palmaris longus

17. Relationship between the m. palmaris longus. and the m.flexor carpi ulnaris

0 They are partially fused

1 They are not fused

18. Origin of the m. adductor pollicis

0 From the ventral surface of the carpal 5-4-3

1 From the internal surface of the distal carpal 5-43

19. Insertion of the m. adductor pollicis. on the prepollex

0 Adjacent to the insertion of the m. abductor pollicis

1 Covers the insertion of the m. abductor pollicis

20. Relationship between the m. adductor pollicis and the m. abductor pollicis

0 Both muscles are independent

1 They are fused

21. Condition of the proximal end of m. lumbricalis brevis indicis

0 Dorsal to the m. adductor pollicis

1 Contiguous to the m. adductor pollicis

22. Origin of the m. lumbricalis brevis Digiti III in relation with the m. tendo superficialis & caput profundum Digiti III

0 Dorsal

1 Laterodorsal

23. Presence of fibers of the tendo superficialis & caput. profundum Digiti III

0 On the external and internal surface of the tendon

1 Only on the internal surface of the tendon

24. Nature of the m. lumbricalis brevis Digiti IV

0 Two slips

1 Only one slip

25. Position of the insertion tendons of the m. lumbricalis longus Digiti IV with respect to the tendo superficialis aponeurosis palmaris prodigiti IV

0 Lateral to the tendo superficialis aponeurosis palmaris prodigiti IV

1 The insertion is via the tendo superficialis aponeurosis palmaris prodigiti IV

26. Origin of the m. lumbricalis longus Digiti V

0 From the aponeurosis palmaris. and TS V

1 Only from TS V

27. Number of insertions of m. lumbricalis longus Digiti V

0 One

1 Two

28. Number of slips that compose the m. lumbricalis brevis Digiti V

0 One

1 Two

Foot

29. Type of insertion of the m. extensor digitorum communis longus

0 Single

1 Multiple

30. Number of insertions of the m. extensor brevis superficialis hallucis

0 Two

1 Three

2 Single

31. Number of insertions of the m. extensor brevis superficialis Digiti II

0 One

1 Two

32. Relationship between the insertions of the m. extensor brevis superficialis Digiti II and the m. extensor digitorum communis longus.

0 Common insertion between m. extensor brevis superficialis Digiti II and the internal slip of the m. extensor digitorum communis longus

1 Inependent insertions

33. Presence of the m. extensor brevis medius hallucis

0 Absent

1 Present

34. Presence of the m. extensor brevis medius Digiti II

0 Absent

1 Present

35. Number of insertion slips of the m. extensor brevis superficialis Digiti III

0 One

1 Two

36. Relationship between the insertions of the mm. extensor brevis superficialis Digiti III and the m. extensor digitorum communis longus.

0 Common insertion between mm. extensor brevis superficialis Digiti III and any slip of m. extensor digitorum communis longus

1 Independent insertions

37. Origin of the m. extensor brevis superficialis Digiti IV

0 By two independent slips

1 By two proximally fused slips

2 By one slip

38. Origin of the m. abductor prehallucis

0 From the internal surface of the aponeurosis plantaris

1 From the distal internal surface of the tarsus

39. Presence of the m. abductor brevis plantaris hallucis

0 Absent

1 Present

40. Type of origin of the m. lumbricalis brevis Digiti III

0 Tendinous

1 Fleshy

41. Relationship between the external slip of m. lumbricalis brevis Digiti III and the internal slip of m. lumbricalis brevis Digiti V

0 Fused only in the origin

1 Fused beyond its origin

Burton characters

Manus

42. Presence of the internal slip of m. lumbricalis brevis Digiti III

0 Absent

1 Present

43. Number of slips of the m. lumbricalis brevis Digiti IV

0 Two

1 Only one, the internal slip

Pes

44. Origin of the tendo superficialis hallucis

0 From the aponeurosis plantaris

1 From the aponeurosis plantaris including some fibers of the m. lumbricalis brevis hallucis

2 From a muscle that originates from the distal tarsal 2-3

45. Origin of the tendo superficialis Digiti II

0 From the aponeurosis plantaris

1 It serves as the insertion tendon of the m. transversus plantae distalis

External characters

46. Skin on venter

0 Areolate

1 Smooth

47. Presence of dorsolateral folds

0 Present

1 Absent

48. Nature of tympannic anulus

0 Complete

1 Incomplete

49. Dorsal view of snout

0 Rounded

1 Subacuminate

2 Acuminate

50. Shape of snout in profile

0 Rounded

1 Truncated

2 Acute

51. Presence of cranial crests

0 Present

1 Absent

52. Width of the upper eyelid in relation with the inter orbital distance

0 Narrower than the inter-orbital distance

1 As wide as the inter-orbital distance

53. Length of fingers I and II

0 First finger shorter than second

1 First and second finger equally or almost equally long

2 First finger longer than second

54. Presence of finger pads or discs

0 Present

1 Absent

55. Presence of skin variations on the ulnar region

0 Present

1 Absent

56. Presence of skin variations on the heel

0 Present

1 Absent

57. Presence of supernumerary plantar tubercles

0 Present

1 Absent

58. Presence of webbed toes

0 Present

1 Absent

59. Presence of toe pads or discs

0 Present

1 Absent

60. Presence of lateral fringes on toes

0 Present

1 Absent

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GALINDO, D. 2004. Filogenia de especies del grupo unistrigatus (sensu Lynch), del genero Eleutherodactylus (Amphibia: Anura: Leptodactylidae) de la cordillera Oriental de Colombia. Trabajo de Grado de Pregrado. Pontificia Universidad Javeriana, Bogota, Colombia. 190 pp.

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LYNCH, J.D. 1975a. A review of the broad-headed eleutherodactyline frogs of South America (Leptodactylidae). Occasional Papers of the Museum of Natural History University of Kansas 38: 1-46.

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LYNCH, J.D. 1989. Intrageneric Relationships of Mainland Eleutherodactylus (Leptodactylidae) I. A Review of the Frogs Assigned to the Eleutherodactylus discoidalis species group.Contributions in Biology and Geology, Milwaukee Public Museum 79: 1-25.

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PYBURN, W.F. & J.D. LYNCH. 1981. Two little-known species of Eleutherodactylus (Amphibia, Leptodactylidae) from the Sierra de la Macarena, Colombia. Proceedings of the Biological Society of Washington 94: 404-412.

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REICHLE, S., S. LOTTERS & I. DE LA RIVA. 2001. A new species of the discoidalis group of Eleutherodactylus (Anura, Leptodactylidae) fron Inner-Andean Dry Valley of Bolivia. Journal of Herpetology 35: 21-26.

RUEDA, J.V. & J.D. LYNCH. 1983. Una nueva especie de Eleutherodactylus (Amphibia: Leptodactylidae) para la Cordillera Oriental de Colombia. Lozania 42: 1-6.

SALGAR, L. 2003. Miologia de manos y pies de Hyla labialis (Peters, 1883) y de Eleutherodactylus bogotensis (Peters, 1883). Trabajo de Grado, Departamento de Biologia, Pontificia Universidad Javeriana Bogota D.C. 258 pp.

SALGAR, L., J.M. HOYOS & A. ACOSTA. 2009. New conditions and intraspecific variations in hand and foot muscles in Pristimantis bogotensis (Peters, 1863) (Amphibia:Anura: Strabomantidae). Zootaxa 2025: 56-68.

Recibido: 12/12/2011

Aceptado: 25/02/2014

JULIO MARIO HOYOS

CATALINA MANTILLA

Pontificia Universidad Javeriana, Departamento de Biologia, Unidad de Ecologia y Sistematica (UNESIS), Apartado 56710, Bogota Colombia. jmhoyos@javeriana.edu.co, dmant010@fiu. edu

DIANA GALINDO

Universidad de los Andes, Departamento de Biologia, Bogota, Colombia. dmgalindo48@uniandes.edu.co

LINA SALGAR

Pontificia Universidad Javeriana, Departamento de Biologia, Unidad de Ecologia y Sistematica (UNESIS), A.A. 56710, Bogota, Colombia. linasalgar@hotmail.com

Table 1. List of species. M: Male.
F: Female.

Specimen    Sex   Species

MUJ 36      M     P. elegans
MUJ 052     F     P. bogotensis
MUJ 465     M     P. bogotensis
MUJ 481     M     P. bogotensis
MUJ 687     F     P. bogotensis
MUJ 23      F     P. nervicus
MUJ 1176    F     P. nervicus
MUJ 1507    F     P. lynchi
MUJ 2036    F     P. lynchi
MUJ 2420    M     P. taeniatus
MUJ 2452    F     P. taeniatus
MUJ 1480    F     P. lynchi
MUJ 1538    M     P. lynchi
ICN 33522   F     P. anolirex
ICN 15444   F     P. anolirex
ICN 26321   M     P. bicolor
ICN 50062   F     P. miyatai
ICN 50065   M     P. miyatai
ICN 50069   F     P. miyatai
ICN 33170   F     P. bacchus
ICN 34232   M     P. bacchus
ICN 33164   M     P. bacchus
ICN 12445   M     P. spilogaster
ICN 12446   F     P. spilogaster
ICN 1932    F     P. lynchi
ICN 1937    F     P. lynchi
ICN 14474   F     P. bicolor
ICN 40579   M     P. frater
ICN 49911   M     P. frater
ICN 15299   M     P. jorgevelosai
ICN 15318   M     P. jorgevelosai
ICN 10107   M     P. prolixodiscus
ICN 10104   F     P. prolixodiscus
ICN 22981   F     P. pugnax
ICN 23187   F     P. pugnax
ICN 21119   M     P. nicefori
ICN 20994   F     P. nicefori
ICN 22949   F     P. tamsiti
ICN 22951   F     P. tamsiti
ICN 23638   F     P. tamsiti
ICN 34233   F     P. merostictus
ICN 34235   F     P. merostictus

Table 2. Data matrix employed in the cladistic
analysis of the LD P unistrigatus species group.

Craugastor raniformis         0000   0010   0000   1010
Pristimantis taeniatus        2000   0001   0000   0010
Pristimantis tamsitti         2000   0001   0010   1010
Pristimantis pugnax           1000   0001   0010   0010
Pristimantis bacchus          1010   1001   0000   0000
Pristimantis miyatai          2010   1001   1100   1010
Pristimantis merostictus      1000   1001   0100   1010
Pristimantis spilogaster      1010   1111   0100   1011
Pristimantis anolirex         2001   1001   0010   0010
Pristimantis jorgevelosai     2001   1001   0010   0010
Pristimantis nicefori         2001   AA10   B010   1000
Pristimantis bicolor          2000   1111   0010   1011
Pristimantis frater           20A1   1101   0001   0100
Pristimantis prolixo discus   2101   1101   0000   0100
Pristimantis bogotensis       2000   0001   0000   0010
Pristimantis elegans          2000   0001   0000   0010
Pristimantis nervicus         2000   000A   0000   0010
Pristimantis lynchi           2000   0001   0100   0010

Craugastor raniformis         0000   0100   0000   0111
Pristimantis taeniatus        0000   0000   0011   1111
Pristimantis tamsitti         1010   1100   0110   0101
Pristimantis pugnax           0000   1001   00C1   0111
Pristimantis bacchus          0000   1000   0010   0110
Pristimantis miyatai          0010   1000   0010   1110
Pristimantis merostictus      0000   0000   0010   0100
Pristimantis spilogaster      0011   0100   0010   0111
Pristimantis anolirex         0000   0001   0011   0111
Pristimantis jorgevelosai     0100   0001   1011   0110
Pristimantis nicefori         0000   0000   0010   1111
Pristimantis bicolor          0000   0111   0111   0111
Pristimantis frater           0000   0001   0010   0011
Pristimantis prolixo discus   0010   0111   101A   0111
Pristimantis bogotensis       0001   1000   0010   0110
Pristimantis elegans          0001   0000   0010   0111
Pristimantis nervicus         0001   0000   0010   1111
Pristimantis lynchi           0001   0100   0010   0111

Craugastor raniformis         10101100111110110100111100011101010000
Pristimantis taeniatus        00011100100100011020110101000000000100
Pristimantis tamsitti         A2010A0120010101A000110100111001001001
Pristimantis pugnax           00010001200001011000110110111000010000
Pristimantis bacchus          11010111200101010000111101010001010100
Pristimantis miyatai          00010001000000010020110100010000000100
Pristimantis merostictus      10010101000001010010110102010000000100
Pristimantis spilogaster      10110101200101010010110001010000000100
Pristimantis anolirex         A00A11010001010Al02Al10001000000000100
Pristimantis jorgevelosai     10010101200101011000110101001000000100
Pristimantis nicefori         10110A01200101010020110000010111010110
Pristimantis bicolor          1001111100010001100021010?010000010100
Pristimantis frater           00010101200000011020110101011000000100
Pristimantis prolixo discus   10A11000000001011020000001210000110100
Pristimantis bogotensis       00111101001000000000110100010001000001
Pristimantis elegans          00011101100001110021110100110001000001
Pristimantis nervicus         10010110100101111020110001210001010100
Pristimantis lynchi           0001110010010001A021110101010000011000

Table 3. Data matrix employed in the cladistic
analysis of the HEA P unistrigatus species group.

Pristimantis frater        1A111010001010000000
                           10000011000101012000
                           01120010101100000100

Pristimantis taeniatus     10000010000001000000
                           00011111000111001010
                           01020010100000000100

Pristimantis tamsitti      10000010010101011011
                           01000101A2010A012011
                           00100010011100101001

Pristimantis pugnax        00000010010001000010
                           10A10111000100012001
                           01100011011100010000

Pristimantis bacchus       01010010000000000010
                           00000110110101112011
                           01000110101000110100

Pristimantis merostictus   00010010100101000000
                           00000100100101010001
                           01010010201000000100

Pristimantis spilogaster   01011110100101101101
                           00000111101101012011
                           01010000101000000100

Pristimantis anolirex      10110010010001000000
                           10010111A00A11010011
                           0A12A000100000000100

Pristimantis nicefori      101AA10B010100000000
                           0000111110110A012011
                           01020000001011110110

Pristimantis bogotensis    10000010000001000110
                           00000110001111010100
                           00000010001000100001

Pristimantis elegans       10000010000001000100
                           00000111000111011001
                           11021010011000100001

Pristimantis nervicus      100000A0000001000100
                           00001111100101101011
                           11120000121000110100

Pristimantis lynchi        10000010100001000101
                           00000111000111001010
                           01A21010101000011000
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Title Annotation:Zoologia-Sistematica
Author:Mario Hoyos, Julio; Mantilla, Catalina; Galindo, Diana
Publication:Caldasia
Date:Jan 1, 2014
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