Rock-dwelling agamids on two continents: Ctenophorus vs Agama, part 1: illustrations

In both Australia and southern Africa, rocky terrain is inhabited by rock-dwelling (rupicolous/saxicoline) lizards in the family Agamidae (https://en.wikipedia.org/wiki/Agamidae).

In Australia, these lizards belong to the genus Ctenophorus (https://en.wikipedia.org/wiki/Ctenophorus).

In southern Africa, the corresponding genus is Agama (https://en.wikipedia.org/wiki/Agama_(lizard)).

In both cases,

  • it is not the genera as such, but instead certain species, that have specialised for a dependence on sheltering in rock crevices, and
  • various congeneric species instead inhabit flat, non-rocky terrain, with various types of vegetation.

In this Post, I focus exclusively on the rock-dwelling species. My aim is to assess the similarities and differences between the two genera in form and function, within the context of rupicolous specialisation.

Two different genera have been recruited on the two landmasses. However, intercontinental similarities in the environment suggest that the lizards should - regardless of phylogeny - be virtually indistinguishable in appearance and biological properties.

In both Ctenophorus and Agama, adult males tend to be colourful to the point of gaudiness. It would be tempting to depict the males here, because they are so photogenic.

However, sexual dimorphism is something of a distraction from the central question of evolutionary convergence/parallelism.

This is because, in many animals, sexual selection tends to produce apparently idiosyncratic results in males. In the case of the lizards examined here, males differ in their conspicuous colouration even from one population to another within a given species (https://static1.squarespace.com/static/514ab09ce4b04d7440ea8696/t/52d8aff4e4b050d4f4e14122/1389932532946/2013McLeanRec.+Aust.+Mus.pdf).

Therefore, the approach I have followed is a scientifically disciplined one, in which I have chosen photos only of adult females and well-grown juveniles.

Unfortunately, the available information on body sizes, given below, does not necessarily specify sex.

CTENOPHORUS DECRESII
total length about 21 cm; maximum 26 cm
snout-vent length mean 7.5-8.2 cm; maximum 9 cm
https://www.sciencedirect.com/science/article/abs/pii/S0003347215004017
https://bmcecolevol.biomedcentral.com/articles/10.1186/s12862-016-0757-2
https://www.inaturalist.org/taxa/31242-Ctenophorus-decresii
juvenile https://www.inaturalist.org/observations/180563545
juvenile https://www.inaturalist.org/observations/173175870
https://www.inaturalist.org/observations/108575781
https://www.inaturalist.org/observations/189946050
https://www.inaturalist.org/observations/70431320
https://www.inaturalist.org/observations/19522971
https://www.inaturalist.org/observations/4925803
?male https://www.inaturalist.org/observations/36461047

CTENOPHORUS FIONNI
total length maximum 28 cm
snout-vent length female 7.8 cm, male 8.7 cm; maximum 9.6 cm
https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1442-9993.2010.02113.x
https://www.publish.csiro.au/ZO/fulltext/ZO20091
https://www.inaturalist.org/taxa/31244-Ctenophorus-fionni
juvenile https://www.inaturalist.org/observations/124213435
https://www.inaturalist.org/observations/153927156
https://www.inaturalist.org/observations/145653566
https://www.inaturalist.org/observations/138221939
https://www.inaturalist.org/observations/138221939
https://www.inaturalist.org/observations/71538380
https://www.inaturalist.org/observations/62616580
https://www.inaturalist.org/observations/54975183
https://www.inaturalist.org/observations/16658359
https://www.inaturalist.org/observations/2391795

CTENOPHORUS GRAAFI
https://www.iucnredlist.org/species/109451364/109451389
https://museum.wa.gov.au/catalogues-beta/digitised-types/herpetology/ctenophorus-caudicinctus-graafi
no observations in iNaturalist

CTENOPHORUS INFANS
snout-vent length maximum 6.7 cm
https://reptile-database.reptarium.cz/species?genus=Ctenophorus&species=infans&search_param=%28%28genus%3D%27E%27%29%29
https://www.inaturalist.org/taxa/889374-Ctenophorus-infans
https://www.inaturalist.org/observations/139180586

CTENOPHORUS MIRRITYANA
total length mean 22 cm, maximum 26 cm
snout-vent length maximum 9.1 cm
https://static1.squarespace.com/static/514ab09ce4b04d7440ea8696/t/52d8aff4e4b050d4f4e14122/1389932532946/2013McLeanRec.+Aust.+Mus.pdf
https://researchoutput.csu.edu.au/en/publications/taxonomic-assessment-of-the-ctenophorus-decresii-complex-reptilia
https://www.inaturalist.org/taxa/554062-Ctenophorus-mirrityana
juvenile https://www.inaturalist.org/observations/41429451
https://www.inaturalist.org/observations/183447920
https://www.inaturalist.org/observations/183446770

CTENOPHORUS MODESTUS
similar to C. decresii
https://www.inaturalist.org/taxa/1250239-Ctenophorus-modestus
https://www.inaturalist.org/observations/188501416
https://www.inaturalist.org/observations/187757947
https://www.inaturalist.org/observations/187207068
https://www.inaturalist.org/observations/176051441
https://www.inaturalist.org/observations/156852092
https://www.inaturalist.org/observations/153920881
https://www.inaturalist.org/observations/138224279
https://www.inaturalist.org/observations/137828734

CTENOPHORUS ORNATUS
total length maximum 29 cm
snout-vent length 8 cm; maximum 9.3 cm
https://www.inaturalist.org/taxa/31227-Ctenophorus-ornatus
http://danielhoops.com/blog/2014/2/16/rock-dragon-group-1
juvenile https://www.inaturalist.org/observations/121705163
https://www.inaturalist.org/observations/188626156
https://www.inaturalist.org/observations/181905372
https://www.inaturalist.org/observations/171728681
https://www.inaturalist.org/observations/170590678
https://www.inaturalist.org/observations/162652008
https://www.inaturalist.org/observations/153317728
https://www.inaturalist.org/observations/148454990
https://www.inaturalist.org/observations/133582040
https://www.inaturalist.org/observations/105144559
https://www.inaturalist.org/observations/91380393
https://www.inaturalist.org/observations/68686041
https://www.inaturalist.org/observations/9607811

CTENOPHORUS RUFESCENS
total length maximum 30 cm
snout-vent length maximum 9.0 cm
https://www.inaturalist.org/taxa/31224-Ctenophorus-rufescens
https://www.inaturalist.org/observations/145710007
https://www.inaturalist.org/observations/168975606
https://arod.com.au/arod/reptilia/Squamata/Agamidae/Ctenophorus/rufescens

CTENOPHORUS SLATERI
snout-vent length maximum 10.0 cm
https://www.inaturalist.org/taxa/889375-Ctenophorus-slateri
juvenile https://www.inaturalist.org/observations/134653940
juvenile https://www.inaturalist.org/observations/95636254
https://www.inaturalist.org/observations/180848666
https://www.inaturalist.org/observations/177966610
https://www.inaturalist.org/observations/177657190
https://www.inaturalist.org/observations/174655377
https://www.inaturalist.org/observations/171513213
https://www.inaturalist.org/observations/171508626
https://www.inaturalist.org/observations/167648701
https://www.inaturalist.org/observations/150391065
https://www.inaturalist.org/observations/122912216
https://www.inaturalist.org/observations/106296140
https://www.inaturalist.org/observations/106296137
https://www.inaturalist.org/observations/95723006
https://www.inaturalist.org/observations/92877557
https://www.inaturalist.org/observations/88920073
https://www.inaturalist.org/observations/34152311
https://www.inaturalist.org/observations/10230867

CTENOPHORUS VADNAPPA
total length maximum 28 cm
snout-vent length maximum 8.6 cm
https://www.inaturalist.org/taxa/31219-Ctenophorus-vadnappa
https://www.inaturalist.org/observations/132799300
https://www.inaturalist.org/observations/145819750

AGAMA ANCHIETAE
total length 15-20 cm, maximum 25 cm
snout-vent length 6-8 cm, maximum 9 cm
https://www.inaturalist.org/taxa/31154-Agama-anchietae
https://www.researchgate.net/publication/335899948_Reproduction_predation_sexual_dimorphism_and_diet_in_Agama_anchietae_Reptilia_Agamidae_from_Namibia
https://namibian.org/nature/reptiles-and-frogs/lizards/anchietas-agama
https://www.inaturalist.org/observations/147847839
https://www.inaturalist.org/observations/118896448
https://www.inaturalist.org/observations/52288001
https://www.inaturalist.org/observations/11244238
https://www.inaturalist.org/observations/11222108
https://www.inaturalist.org/observations/119357
Feminine colouration (females in breeding condition):
https://www.flickr.com/photos/martin_hingst/12259309126 and https://www.inaturalist.org/observations/63882192 and https://www.inaturalist.org/observations/10819713 and https://www.inaturalist.org/observations/172317220 and https://www.inaturalist.org/observations/146603496

AGAMA ATRA
total length up to about 25 cm; 20-25 cm, maximum 32 cm
snout-vent length 7-12 cm; female 9.2 cm, male 12.5 cm
snout-vent length maximum females 10 cm, males 10.9 cm, except in Namaqualand, where females 11 cm, males 13.9 cm (https://web.archive.org/web/20160304185710/http://academic.sun.ac.za/capeherp/cederberg/rockagama.htm)
https://onlinelibrary.wiley.com/doi/abs/10.1111/mec.15901
https://scholar.sun.ac.za/items/d0a94bd8-bfcc-402b-83ce-07d944039bd8
https://onlinelibrary.wiley.com/doi/abs/10.1046/j.0962-1083.2001.01458.x
https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1365-2699.2008.01988.x
https://www.inaturalist.org/taxa/31150-Agama-atra
https://www.inaturalist.org/observations/165155507
https://www.inaturalist.org/observations/187815636
https://www.inaturalist.org/observations/187216307
https://www.inaturalist.org/observations/185887296
https://www.inaturalist.org/observations/184233155
https://www.inaturalist.org/observations/182839544
https://www.inaturalist.org/observations/182312800
https://www.inaturalist.org/observations/181591198
https://www.inaturalist.org/observations/179699655
https://www.inaturalist.org/observations/178950565
https://www.inaturalist.org/observations/177341792
https://www.inaturalist.org/observations/170532824
https://www.inaturalist.org/observations/156405081
https://www.inaturalist.org/observations/152264717
https://www.inaturalist.org/observations/152248008
https://www.inaturalist.org/observations/149743803
https://www.inaturalist.org/observations/148914255
https://www.inaturalist.org/observations/144145227
https://www.inaturalist.org/observations/142865290
https://www.inaturalist.org/observations/141354843
https://www.inaturalist.org/observations/140246454
Feminine colouration (females in breeding condition):
https://www.inaturalist.org/observations/105045352 and https://www.inaturalist.org/observations/147245252 and https://www.inaturalist.org/observations/145560986 and https://www.inaturalist.org/observations/135279556 and https://www.inaturalist.org/observations/126623121

AGAMA KIRKII
total length 18-25 cm, maximum 34 cm
female snout-vent length 7.7-8.1 cm (https://www.gbif.org/species/5226328)
https://www.inaturalist.org/taxa/31136-Agama-kirkii
https://www.inaturalist.org/observations/114206705
https://www.inaturalist.org/observations/103148911
https://www.inaturalist.org/observations/46875905
https://www.inaturalist.org/observations/41675619
https://www.inaturalist.org/observations/40872615
https://www.inaturalist.org/observations/18086076
Feminine colouration (females in breeding condition):
https://www.inaturalist.org/observations/10005155 and https://www.inaturalist.org/observations/9099154 and https://www.inaturalist.org/observations/143078211 and https://www.inaturalist.org/observations/25819395

AGAMA KNOBELI
https://www.inaturalist.org/taxa/539228-Agama-knobeli
https://reptile-database.reptarium.cz/species?genus=Agama&species=knobeli
https://www.inaturalist.org/observations/186980177
https://www.inaturalist.org/observations/1050044

AGAMA PLANICEPS
total 25-30 cm, maximum 33 cm
snout-vent length 8-10 cm, maximum 11.2 cm
https://www.inaturalist.org/taxa/31127-Agama-planiceps
https://namibian.org/nature/reptiles-and-frogs/lizards/namibian-rock-agama
https://www.tyroneping.co.za/lizards/agama-planiceps/
https://animalia.bio/agama-planiceps
https://www.inaturalist.org/observations/11244125
https://www.inaturalist.org/observations/9032266
https://www.inaturalist.org/observations/11245118
https://www.inaturalist.org/observations/11222143
https://www.inaturalist.org/observations/11349047
https://www.inaturalist.org/observations/188529479
https://www.inaturalist.org/observations/187623714
https://www.inaturalist.org/observations/183642841
https://www.inaturalist.org/observations/180470859
https://www.inaturalist.org/observations/177496268
https://www.inaturalist.org/observations/161592415
https://www.inaturalist.org/observations/149753925
https://www.inaturalist.org/observations/147143136
https://www.inaturalist.org/observations/146407326
https://www.inaturalist.org/observations/144961896
https://www.inaturalist.org/observations/142677426
https://www.inaturalist.org/observations/139989413
https://www.inaturalist.org/observations/136805245
https://www.inaturalist.org/observations/136011887
https://www.inaturalist.org/observations/126033865
https://www.inaturalist.org/observations/125674252
https://www.inaturalist.org/observations/120644539
https://www.inaturalist.org/observations/112171810
https://www.inaturalist.org/observations/112032010
https://www.inaturalist.org/observations/107382194
https://www.inaturalist.org/observations/103038168
https://www.inaturalist.org/observations/101460621
https://www.inaturalist.org/observations/92664684
https://www.inaturalist.org/observations/69873672
https://www.inaturalist.org/observations/59813063
https://www.inaturalist.org/observations/58768113
https://www.inaturalist.org/observations/56241844
https://www.inaturalist.org/observations/48470549
https://www.inaturalist.org/observations/47359777
https://www.inaturalist.org/observations/34609853
https://www.inaturalist.org/observations/14929173
https://www.inaturalist.org/observations/13694859
https://www.inaturalist.org/observations/14378017

to be continued in https://www.inaturalist.org/journal/milewski/86326-rock-dwelling-agamids-on-two-continents-ctenophorus-vs-agama-part-2#...

Publicado el 29 de octubre de 2023 por milewski milewski

Comentarios

https://www.researchgate.net/publication/335899948_Reproduction_predation_sexual_dimorphism_and_diet_in_Agama_anchietae_Reptilia_Agamidae_from_Namibia:

Agama anchietae differs from A. planiceps in being smaller-bodied, less sexually dimorphic, solitary as opposed to social, not polygynous to the same degree, and inconspicuous in colouration.

The diets are similar (ants, termites, and other insects), except that anchietae does not eat beetles as frequently as does planiceps.

Agama anchietae lays more eggs per clutch than does A. planiceps, compensating by the eggs being relatively small.

I infer that the two species coexist at the hectare scale, despite belonging to the same genus, and having similar diets.

It is not habitat or diet that separate them, but instead social/sexual/reproductive strategy, and anti-predator strategy. Agama planiceps, unlike A. anchietae, is large, conspicuous, and particularly social/polygynous.

Publicado por milewski hace 8 meses
Publicado por milewski hace 8 meses

Information on the Web, for snout-vent lengths in rupicolous spp. of Agama, is limited. This hinders comparison with Ctenophorus, in which snout-vent length is the usually-reported measure of body size (the rock-dwelling spp. have snout-vent lengths of about 7-10 cm).

Agama atra is known to have snout-vent length about 9 cm in females, and about 12.5 cm in males (Bruton, 1977).

Agama agama is not specialised for rock-dwelling (Harris, 1964, https://www.amazon.com/Life-Rainbow-Lizard-V-Harris/dp/0090721403 and https://www.biblio.com/book/life-rainbow-lizard-harris-v/d/1288437620).

Enge et al. (2004, https://www.jstor.org/stable/24321176) state that adult females of A. agama have snout-vent length 9.4 cm or more. This indicates that A. agama exceeds A, atra in body size.

A study at the delta of the Niger River (https://abjournals.org/african-journal-of-biology-and-medical-research-ajbmr/wp-content/uploads/sites/17/journal/published_paper/volume-3/issue-1/AJBMR_IgtNPMp9.pdf) further documents A. agama.

However, a) both adults and juveniles were sampled (which is not stated in the Methods but is evident from the ranges of values of body size), and b) what is stated to be 'snout-vent length' seems actually to be total length.

The authors sampled of 151 individuals of the two sexes combined, in February to May.

They do not state whether females were gravid or not. However, I infer that they were not at this time, because sampling was done late in the dry season.

The findings, for A. agama, are:

Females (not necessarily adult):
Mean body mass 20-22 g
Total length ('misreported as 'snout vent length') 18-20.5 cm

Males (not necessarily mature):
Mean body mass 48-51 g
Total length (misreported as 'snout vent length') 26.5 cm

I infer from these poorly-presented data that, in A. agama, adult females have total length about 28 cm and body mass about 45 g, whereas mature males have total length about 32 cm and body mass about 70 g.

Agama planiceps is similar to A. agama (which is large-bodied for its genus) in body size. Thus, the body mass of adult females of A. planiceps is double that typical for adult females of rock-dwelling spp. of Ctenophorus.

I infer that A. planiceps is not only more social/gregarious/colonial than any counterpart in Australia, but also larger-bodied - even if we correct for the extreme sexual dimorphism in A. planiceps

Publicado por milewski hace 8 meses

Agama agama are big animals, 20cm sounds right for the SVL of an adult.
Original descriptions and the old school expedition reports have information such as size, but not so much nowadays. You will need to dig a bit. Otherwise check Branch 1998 for a summary of southern African agamid sizes.

Publicado por alexanderr hace 8 meses

https://onlinelibrary.wiley.com/doi/epdf/10.1111/j.1600-0587.1999.tb00503.x

This reference gives asymptotic snout-vent lengths as Agama impalearis = bibronii (https://www.inaturalist.org/taxa/1367368-Agama-bibronii) grows to adulthood.

Near Marrakech (https://en.wikipedia.org/wiki/Marrakesh) in Morocco, asymptotic snout-vent length has mean values of a) 12.5 cm in males, and b) 11.4 cm in females.

Sexual maturity occurs at mean snout-vent length of a) 9.1 cm in males, and b) 8.8 cm in females.

Within Morocco, the asymptotic snout-vent lengths of males vary from means of about 13 cm in the southwest of the country to means of about 12 cm in the northeast of the country.

These are the most reliable data that I have found, so far, for snout-vent lengths in any rupicolous spp. of Agama. How do they compare with Ctenophorus?

(An obvious caveat is that Agama impalearis = bibronii occurs in North Africa, whereas my study is in southern Africa.)

Well, snout-vent lengths for adults of Ctenophorus spp. are about 7-10 cm.

This indicates that body size is considerably greater in rupicolous Agama (snout-vent lengths > 11.5 cm) than in rupicolous Ctenophorus (snout-vent lengths < 10 cm).

Publicado por milewski hace 8 meses

@alexanderr

Do you by any chance have access to this article?

https://www.tandfonline.com/doi/abs/10.1080/15627020.2000.11657095

Publicado por milewski hace 8 meses

https://www.researchgate.net/profile/El-Hassan-El-Mouden/publication/249645371_Sexual_dimorphism_reproductive_and_fat_body_cycles_in_Bibron%27s_agama_Agama_impalearis_Boettger_1874Sauria_Agamidae/links/55b8ad7808ae9289a08dd62b/Sexual-dimorphism-reproductive-and-fat-body-cycles-in-Bibrons-agama-Agama-impalearis-Boettger-1874Sauria-Agamidae.pdf

AGAMA BIBRONII = IMPALEARIS

Central Jbilet mountains, Morocco

"mature males ranged from 91-134 mm SVL and from 25.5-100 g in mass, and mature females ranged from 88-125 mm SVL and from 27-85 g in mass (excluding clutch mass in gravid females). The mean SVL was of 113.09...mm and 105.77...mm, respectively for males and females corresponding to average body masses of 62.60...g and 51.78...g.

I note that body mass of adult females in this species exceeds that of any of the southern African rock-dwelling congeners.

Publicado por milewski hace 8 meses

Bruton (1977, https://www.tandfonline.com/doi/abs/10.1080/00445096.1977.11447557) shows the following snout-vent lengths for Agama atra in Eastern Cape province:

adult female: 9.2 cm

adult male: 12.5 cm

Publicado por milewski hace 8 meses

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