Let us first start with an instructive presentation by the well known
herpetologist Stephen Spawls.
City College Norwich biology and chemistry lecturer, Mr Spawls, from Spixworth,
grew up in Kenya and it was here that he developed his enduring interest as a
herpetologist. He has had a lifelong interest in African wildlife, and a
particular passion for snakes and chameleons. For the books he wrote about
African herpetology,
click here.
Click on the picture or
here to see the presentation, or click to download a
Powerpoint
Presentation.
Classification schemes are dependant on the actuality of papers on which they are based. A full classification scheme can be found on the webiste of Animal Diversity Web, but this is already outdated.
Vidal e.a. 2007 "investigated the higher-level relationships of caenophidian snakes with seven nuclear protein-coding genes and obtained a well-supported topology. Accordingly, some adjustments to the current classification of Caenophidiaare made to better reflect the relationships of the groups. The phylogeny also indicates that, ancestrally, caenophidian snakes areAsian and nocturnal in origin, although living species occur on nearly all continents and are ecologically diverse." The for our group of snakes relevant part of their taxonomic classification scheme is:
Superfamily Elapoidea Boie, 1827
Family Elapidae Boie, 1827
(Subfamilies: Elapinae Boie, 1827 and Hydrophiinae Fitzinger, 1843)
Family Lamprophiidae Fitzinger, 1843
(Subfamilies: Psammophiinae Bonaparte, 1845, Atractaspidinae Günther, 1858,
“Lamprophiinae” Fitzinger, 1843 and “Pseudoxyrhophiinae” Dowling, 1975)
In Vidal e.a. (2008) the
classification is given more in detail:
" The Elapoidea includes the Elapidae and a large (~60 genera, 280 sp.) and
mostly African (including Madagascar) radiation termed Lamprophiidae by Vidal et
al. (2007), that includes at least four major groups: the psammophiines,
atractaspidines, lamprophiines and pseudoxyrhophiines." And: "The lamprophiids
(~60 genera, 280 sp.) include four major groups: the psammophiines (~7 genera,
42 sp.), atractaspidines (~12 genera, 70 sp.), lamprophiines (~ 19 genera, 88 sp.)
and pseudoxyrhophiines (~20 genera, 80 sp.) (Vidal 2002; Vidal & Hedges 2002a;
Vidal et al. 2007). Following Vidal et al. (2007, 2009), Fry et al. (2008), and
Vonk et al. (2008), we treat these groups as subfamilies of the African
lamprophiid radiation, although others have afforded them family status,
including Lamprophiidae, Psammophiidae, Pseudoxyrhophiidae (Kelly et al. 2008)
and Atractaspididae (Branch 1998; Zaher 1999; Shine et al. 2006), while
additional families, Prosymnidae (genus Prosymna) and Pseudaspididae
(genera Pseudaspis and Pythonodipsas), have recently been proposed
(Kelly et al. in press).
Scheme from Vidal a.o. 2007:
First, Pyron e.a. define crown-group Colubroidea to consist of the extant families Colubridae, Elapidae, Homalopsidae, Lamprophiidae, Pareatidae, Viperidae, and Xenodermatidae. The result for the psammophiids is:
Superfamily Colubroidea. Family Lamprophiidae, Subfamily Psammophiinae
In their view the correct term for the family is also Psammophiinae.
A recent paper is Christopher M.. Kelly, Nigel P. Barker, Martin H. Villet, Donald G. Broadley and William R. Bran, The snake family Psammophiidae (Reptilia: Serpentes): Phylogenetics and species delimitation in the African sand snakes (Psammophis Boie, 1825) and allied genera. (2008). They describe the family as follows:
"Psammophiidae Bourgeois, 1968 occurs throughout Africa, the Middle East, Madagascar, southern Europe and south-central Asia, and currently includes eight genera and about 50 extant species (Table 1), of which around 30 belong to the type genus Psammophis Boie, 1825. Other genera include Dipsina Jan, 1863 (one species), Dromophis Peters, 1869 (two species), Hemirhagerrhis Boettger, 1893 (four species), Malpolon Fitzinger, 1826 (three species), Mimophis Günther, 1868 (one species), Psammophylax Fitzinger, 1843 (four species), and Rhamphiophis Peters, 1854 (five species)."
Concerning the place of former Rhamphiophis acutus within the
family, they conclude:
"In clade B, the strong support for a sister relationship
between
Psammophylax and
R. acutus
renders the genus
Rhamphiophis
diphyletic. Broadley (1971) provides a review and
taxonomic history of
R. acutus,
noting that in several respects (dentition and colouration) this taxon forms a
link between
Rhamphiophis and
Psammophylax.
The most distinctive synapomorphy of
Rhamphiophis
is its shortened and reinforced skull (adaptations for
digging), a characteristic that is less pronounced in
R. acutus
([Broadley, 1971] and [Chirio and Ineich, 1991]). It is clear
that this character is homoplasious, and we transfer the three subspecies of
R. acutus
to the genus
Psammophylax,
i.e.
Psammophylax acutus acutus (Günther, 1888) (comb. nov.),
Psammophylax
acutus jappi (Broadley, 1971) (comb. nov.) and
Psammophylax acutus
togoensis (Matschie, 1893) (comb. nov.). "
The consequence for the dentition description of Psammophylax is:
"This change increases the dental variation in
Psammophylax
and necessitates the following modification to the
diagnosis provided by Broadley (1977b): ‘The maxilla bears 9–13 subequal teeth,
separated by a diastema from two enlarged grooved fangs on the posterior end of
the bone; palatine teeth 7–11; pterygoid 9–23; dentary 15–24.’"
Concerning the place of Dromophis within the family, they conclude:
"Dromophis praeornatus
and
Dromophis lineatus
occupy divergent and
well-supported positions nested deeply within
Psammophis, in
clades 5 and 7 respectively (Fig. 1). We therefore relegate the genus
Dromophis
to the synonymy of
Psammophis,
forming Psammophis praeornatus
(Schlegel, 1837) (comb.
nov) and Psammophis lineatus
(Dumèril & Bibron, 1854) (comb.
nov.). "
The consequence for the dentition description of Psammophis is:
"The generic diagnosis provided for
Psammophis
by Broadley (1983) should thus be modified as follows: ‘Maxillary
teeth usually 10–16, 3–5 small teeth anteriorly, followed after a distinct
interspace by two much enlarged recurved and fang-like teeth (below anterior
border of eye), which are followed, after a second interspace, by 3–7 small
teeth and then two strongly enlarged grooved fangs (below the posterior border
of the eye). In
P. praeornatus and
P. lineatus
(formerly assigned to the genus
Dromophis)
there is a continuous series of 10–16 small teeth (median longest) preceding the
large grooved fangs below the posterior border of the eye. Anterior mandibular
teeth strongly enlarged.’ "
Within the genus, Psammophis sibilans is the most commonly cited species and the most problematic. For a long time, P. sibilans was considered a species widespread throughout most of sub-Saharan Africa with variable coloration. This incredible phenotypic variation across its wide distribution range led to the description of several “races”, varieties and subspecies, some of them were shown to have no support, while others are currently considered full species. However, this situation often leads to misidentifications and the excessive synonymization, resulting in a highly confusing taxonomic history. (Parrinha 2020)
As for this sibilans-complex (P. sibilans, P. phillipsi, P. sudanensis, P. mossambicus, P. leopardinus, P. brevirostris, P. rukwae, P. subtaeniatus and P. orientalis), I advise you to read the recent (2020) study by Parrinha in which he sets a first step towards a taxonomic revision.
Colour and markings
Parrinha’s results produced from data associated with color pattern (presence and type of head markings, dorsolateral stripes, and ventral lines) show considerable variation in all species within this group. In P. sibilans there are usually distinct head markings and dorsolateral stripes, however, completely uniform specimens are not rare. Specimens with a uniform ventrum and continuous ventral stripes occur in similar proportions, but discontinuous lines or series of dots are uncommon. In P. mossambicus the top of the head is almost always uniform. Even though the results show the occurrence of dorsolateral and ventral stripes in P. mossambicus, these specimens are mostly restricted to Central Africa, so that outside this area P. mossambicus is almost always dorsally uniform and without continuous ventral lines. P. sudanensis shows the least variation in color pattern, with most specimens displaying well-defined head markings, as well as dorsal and ventral stripes. Parrinha did not find differences in coloration between East African P. rukwae and P. cf. rukwae from West Africa. Both taxa may be striped or uniform, with or without head markings, and more than 50% of specimens of both species display thin ventral lines.
Philippe Geniez, Alexandre Cluchier & Cornelius C. De Haan, 2006 describe
the Psammophiinae in the following way.
Psammophines are a colubrid tribe or subfamily of mostly African snakes,
consisting of 8 genera and about 44 species. These are characterized, in males, by their tiny hemipenes, quasi filiform
and 3-4 subcaudals short (BOGERT 1940)
and, in both sexes, by their valvular nostril enabling “self-rubbing” (DE HAAN
2003, DE HAAN & CLUCHIER 2005). Moreover, lack of significant sexual dimorphism
in tail length seems to be generalized in the whole tribe, anyway for sure in
Malpolon monspessulanus (DE HAAN 1999).
Picture from Bogert 1940.
Sexing these snakes is very difficult. One safe methode is taking a close look at the anal part of shed skins. See the pictures of the sheddings of some males on this site.
Broadley
knew it back in 1959: "Sexual dimorphism. The sexes cannot be separated
on scale counts. Sexing is made difficult by the small size of the
hemipenes. The everted hemipenes of a 1466 mm. Essexvale males were only
12 mm. in length and 2 mm. in diameter.
Picture thanks to Cornélius de Haan
Dentition of Psammophiids, picture from Bogert 1940.
According to Cottone & Bauer 2009, sand snakes of the genus Psammophis consist of ~30 species in which species boundaries and taxonomic allocations have been widely disputed. Psammophis species are often the most common and conspicuous snakes throughout their ranges in Africa and Palearctic Asia. Unlike most snakes, psammophiids in tropical and temperate regions display little to no sexual size dimorphisme (SSD), exhibit little intrageneric variation in diet, and generally have low fecundities.
I quote a short discussion on the timeline of Nicole Tam on Facebook from January 2019:
"Martin Habecker WC ones are readily available and are extremely
hardy captives. Minus the P. tanganicus, I have had over a dozen Psammophis
species over the last 12yrs and never had an issue with acclimation with them.
I'd go with Nicole Tam's offer though when she has some available but babies can
be very tricky to care for. I sent my wrongly assumed male P. mossambicus to
Myke Clarkson who had a wrongly assumed female (funny how things work out with
Psammophis isn't it) and I got two offspring from him. I had to hand feed them
Anole tails for a while, I believe Ton Steehouder has the pics up on the
Psammophid site, so it was a little tricky. Adults never refused a meal though!
Nicole Tam
Nicole Tam Martin Habecker that's what I'm worried about too, the care of the
babies. I don't think I'm up for some serious breeding efforts with mine yet,
especially considering recent events on top of these rostratus (they're fine,
just a lot of work).
Rupert Cole, Fascination Herp looks like they have some WC ones now actually,
that's where I got my group of four from.
Martin Habecker
Martin Habecker I made a corny video a few years back feeding some of my African
colubrid collection for fun primarily, but also to show that most of these then
termed "difficult" species were actually quite amicable to F/T rodent prey
items. Ignore the music, I am a silly person https://www.youtube.com/watch?v=H2Ihhh2qdsQ
youtube.com, .
African colubrid feeding2
Nicole Tam
Nicole Tam Delightfully corny music!
Rupert Cole
Rupert Cole As soon as I get home I am watching it.
Martin Habecker
Martin Habecker Well on the dl, if you guys want them I can get you much better
deals on them in the future. I pay quite a bit less for WC imports but I am not
getting any in until the weather warms up a bit.
Nicole Tam
Nicole Tam Awesome! Thank you Martin, I'll keep that in mind.
Rupert Cole
Rupert Cole Thanks Martin Habecker! I would love to get them from you.
Martin Habecker
Martin Habecker Anytime, gotta keep the sweet Psammophis a secret though too
many people are asking questions of late
Rupert Cole
Rupert Cole
Martin Habecker
Martin Habecker As an addendum, the M. poensis is actually Gonionotophis (Mehelya)
crossi.
Nicole Tam
Nicole Tam Agreed...that's odd. Wonder why such a big surge in interest. But, my
lips are sealed.
Martin Habecker
Martin Habecker Not sure, I think that Andrew Durso's blog increased interest in
the small but fanatical whipsnake and oddball colubrid crowd, that our
collective knowledge of herpetoculture has expanded to include lesser known
snakes that were previously sold as feed…Meer bekijken
Martin Habecker
Martin Habecker I kept seeing them pop up on import lists for next to nothing,
did a little research found DeHaan's paper on them as well as the Psammophis
website and purchased a group of 2 P. tanganicus, 3 P. sibilans, and 1 P.
mossambicus for about $120shipped next time I saw them available back in 2007
and have raved about them ever since.
Nicole Tam
Nicole Tam Martin Habecker Funny how interest spreads around, right? Reptiles
Magazine was my starting point for Psammophiinae, right at the first article on
Rhamphiophis rostratus. Fast forward more than 10 years and here I am, getting
R. rostratus to make a 4.2Funny how interest spreads around, right? Reptiles
Magazine was my starting point for Psammophiinae, right at the first article on
Rhamphiophis rostratus. Fast forward more than 10 years and here I am, getting
R. rostratus to make a 4.2.1 group (and breeding them too! Younger me would have
been mindblown). Then, joining Ton's group and slowly getting around to talking
with people like you, Ton, and Cornelius.
And for Malpolon especially, learning about them roughly 5 years ago from my
husband as he'd see them and Dolichophis jugularis in Lebanon. I would have
never expected to even have such awesome animals.
It's been a hell of a ride and I don't regret it one bit. There's always
something fun to learn about Psammophiinae. And, of course, dragging people like
Rupert Cole in like the bad influence I am. "
A wonderful website about the herpetofauna of Ethiopia and adjacent territory
can be found on
http://www.reptiles-of-ethiopia-and-eritrea.com.
The author is Vladimir Trailin. Many
psammophiid snakes are treated there, with beautiful pictures.
