329
Diagnoses for the Newly Established Taxa
Suborder Tendrina subord. nov.
Diagnosis as in family.
Superfamily Tendroidea Vigneaux, 1949
Diagnosis as in family.
Family Tendridae Vigneaux, 1949
Diagnosis . Colony encrusting, uniserial to loosely pluriserial and multiserial. Autozooids of malacostegan-grade
structure, with well-developed smooth gymnocyst and large
oval opesia. Mural spines normally present, articulated and
non-articulated. No avicularia. Simultaneous incubation of
several embryos of same age in acanthostegal brood chamber
formed by mural spines of either autozooid ( Tendra ) or distal
kenozooid ( Heteroecium ). In latter case it fuses with eggproducing autozooidal polymorph to form cormidial brood
complex. Intertentacular organ may be present (recorded in
Tendra ). Lateral septula multiporous. Ancestrula
autozooidal.
Time range : Recent.
Remarks . The structure of the acanthostegal brood chambers indicates that they (and the non-feeding larva) must
have originated independently of the ovicells and larvae in
Flustrina. The most recent ancestor of Tendridae was a malacostegine cheilostome with an intertentacular organ and nonarticulated mural spines. Multiplication and compaction of
these spines was a necessary fi rst step in the origin of the
protective ‘roof’ of the brood chamber.
Suborder Thalamoporellina subord. nov.
Diagnosis as in superfamily.
Superfamily Thalamoporelloidea Levinsen, 1902
Diagnosis . Colony encrusting or erect. Zooids with
depressed porous cryptocyst which generally reaches the
opercular area; if so, one or a pair of large opesiules and a
polypide tube are present, and there is a small opesia coincident with the orifi ce. Internal free spicules usual. Avicularia,
when present, vicarious or subvicarious/interzooidal.
Embryos brooded in ovicells (Thalamoporellidae) or internal
brood sacs (Steginoporellidae). Ovicells, when present,
hyperstomial, cleithral, simultaneously incubating two to
several embryos of different ages. Ooecium with median
suture, formed from maternal autozooid. Large intertentacular
organ may be present. Lateral septula multiporous or biporous. Ancestrula autozooidal or kenozooidal.
Time range : Lutetian (Early-Middle Eocene) – Recent.
Remarks . Among both Thalamoporellidae and
Steginoporellidae, Thalamoporella is exceptional in possessing ovicells whose structure, placement and inception indicate that they (and the non-feeding larva) must have
originated independently of Flustrina. Thalamoporella and
Steginoporella , both known from the Lutetian evidently had
a malacostegine ancestor. Thalamoporella evolved by independently acquiring an extensive cryptocyst and bilobate
ovicells formed by the extension and fusion of a pair of frontal tubercles. The evolution of brooding was accompanied by
a shift from larval planktotrophy to lecithotrophy. Ovicells
were substituted by internal brood sacs in steginoporellids
and in other thalamoporellids. ‘Marsupial’ ovicell-like structures situated proximal to the autozooidal orifi ce of the
Recent Marsupioporella , and the reduction of the cryptocyst
in Hesychoxenia were apparently later developments. Note
that the Cretaceous genus Dimorphomicropora provisionally
included by some in the Steginoporellidae, was probably an
unrelated thalamoporelline homeomorph.
Suborder Belluloporina subord. nov.
Diagnosis as in family.
Superfamily Belluloporoidea superfam. nov.
Diagnosis as in family.
Family Belluloporidae fam. nov.
Diagnosis . Colony encrusting, multiserial. Autozooids of
cribrimorph-grade organization, with very narrow smooth
gymnocyst and frontal membranous wall overarched by costae. Each costa a kenozooid with a long strip of hypostegal
coelom confl uent with visceral coelom of autozooid via a
communication pore with a cuticular annulus. Costae joined
to their neighbours via short lateral fusions, leaving rows of
bean-shaped lacunae between fusions. Articulated oral
spines present. Pedunculate adventitious avicularia on either
side of zooidal aperture. Ovicell hyperstomial, cleithral,
formed by fused costae of the distal kenozooid that has
membranous frontal wall serving as ovicell fl oor. Basal pore
chambers present. Ancestrula tatiform with mural spines surrounding frontal membrane.
Time range : Pleistocene – Recent.
Remarks . The kenozooidal nature of the ooecial costae
indicates that these brood chambers must have originated
independently of Flustrina. While the larva of Bellulopora
has never been described, the suggestion that it is non- feeding
is plausible. The putative ancestor of Bellulopora would have
had a malacostegan-grade organization with numerous mural
kenozooidal spines. Their fusion led to an independent origin
of the spinocyst (and costate ooecia) in this genus. The example of Belluloporidae indicates that cheilostome spines may
originally have been zooid polymorphs. It is rather probable
that similar cheilostomes (both malacostegans and ‘cribrimorphs’) with kenozooidal spines should be more widespread, but their recognition in the fossil record is problematic
since the organic annulus that is characteristic of conventional
communication pores is not preserved.
Superfamily Monoporelloidea Hincks, 1882
Diagnosis . Colony encrusting (typically multiserial,
sometimes uniserial or pluriserial) or erect (bilaminar-folded
or dichotomously branching with fl exible nodes). Zooids
with opesiules and pores in an extensive cryptocyst, forming
a small opesia coincident with the orifi ce. Gymnocystal rim
Appendices
Diagnoses for the Newly Established Taxa
Suborder Tendrina subord. nov.
Diagnosis as in family.
Superfamily Tendroidea Vigneaux, 1949
Diagnosis as in family.
Family Tendridae Vigneaux, 1949
Diagnosis . Colony encrusting, uniserial to loosely pluriserial and multiserial. Autozooids of malacostegan-grade
structure, with well-developed smooth gymnocyst and large
oval opesia. Mural spines normally present, articulated and
non-articulated. No avicularia. Simultaneous incubation of
several embryos of same age in acanthostegal brood chamber
formed by mural spines of either autozooid ( Tendra ) or distal
kenozooid ( Heteroecium ). In latter case it fuses with eggproducing autozooidal polymorph to form cormidial brood
complex. Intertentacular organ may be present (recorded in
Tendra ). Lateral septula multiporous. Ancestrula
autozooidal.
Time range : Recent.
Remarks . The structure of the acanthostegal brood chambers indicates that they (and the non-feeding larva) must
have originated independently of the ovicells and larvae in
Flustrina. The most recent ancestor of Tendridae was a malacostegine cheilostome with an intertentacular organ and nonarticulated mural spines. Multiplication and compaction of
these spines was a necessary fi rst step in the origin of the
protective ‘roof’ of the brood chamber.
Suborder Thalamoporellina subord. nov.
Diagnosis as in superfamily.
Superfamily Thalamoporelloidea Levinsen, 1902
Diagnosis . Colony encrusting or erect. Zooids with
depressed porous cryptocyst which generally reaches the
opercular area; if so, one or a pair of large opesiules and a
polypide tube are present, and there is a small opesia coincident with the orifi ce. Internal free spicules usual. Avicularia,
when present, vicarious or subvicarious/interzooidal.
Embryos brooded in ovicells (Thalamoporellidae) or internal
brood sacs (Steginoporellidae). Ovicells, when present,
hyperstomial, cleithral, simultaneously incubating two to
several embryos of different ages. Ooecium with median
suture, formed from maternal autozooid. Large intertentacular
organ may be present. Lateral septula multiporous or biporous. Ancestrula autozooidal or kenozooidal.
Time range : Lutetian (Early-Middle Eocene) – Recent.
Remarks . Among both Thalamoporellidae and
Steginoporellidae, Thalamoporella is exceptional in possessing ovicells whose structure, placement and inception indicate that they (and the non-feeding larva) must have
originated independently of Flustrina. Thalamoporella and
Steginoporella , both known from the Lutetian evidently had
a malacostegine ancestor. Thalamoporella evolved by independently acquiring an extensive cryptocyst and bilobate
ovicells formed by the extension and fusion of a pair of frontal tubercles. The evolution of brooding was accompanied by
a shift from larval planktotrophy to lecithotrophy. Ovicells
were substituted by internal brood sacs in steginoporellids
and in other thalamoporellids. ‘Marsupial’ ovicell-like structures situated proximal to the autozooidal orifi ce of the
Recent Marsupioporella , and the reduction of the cryptocyst
in Hesychoxenia were apparently later developments. Note
that the Cretaceous genus Dimorphomicropora provisionally
included by some in the Steginoporellidae, was probably an
unrelated thalamoporelline homeomorph.
Suborder Belluloporina subord. nov.
Diagnosis as in family.
Superfamily Belluloporoidea superfam. nov.
Diagnosis as in family.
Family Belluloporidae fam. nov.
Diagnosis . Colony encrusting, multiserial. Autozooids of
cribrimorph-grade organization, with very narrow smooth
gymnocyst and frontal membranous wall overarched by costae. Each costa a kenozooid with a long strip of hypostegal
coelom confl uent with visceral coelom of autozooid via a
communication pore with a cuticular annulus. Costae joined
to their neighbours via short lateral fusions, leaving rows of
bean-shaped lacunae between fusions. Articulated oral
spines present. Pedunculate adventitious avicularia on either
side of zooidal aperture. Ovicell hyperstomial, cleithral,
formed by fused costae of the distal kenozooid that has
membranous frontal wall serving as ovicell fl oor. Basal pore
chambers present. Ancestrula tatiform with mural spines surrounding frontal membrane.
Time range : Pleistocene – Recent.
Remarks . The kenozooidal nature of the ooecial costae
indicates that these brood chambers must have originated
independently of Flustrina. While the larva of Bellulopora
has never been described, the suggestion that it is non- feeding
is plausible. The putative ancestor of Bellulopora would have
had a malacostegan-grade organization with numerous mural
kenozooidal spines. Their fusion led to an independent origin
of the spinocyst (and costate ooecia) in this genus. The example of Belluloporidae indicates that cheilostome spines may
originally have been zooid polymorphs. It is rather probable
that similar cheilostomes (both malacostegans and ‘cribrimorphs’) with kenozooidal spines should be more widespread, but their recognition in the fossil record is problematic
since the organic annulus that is characteristic of conventional
communication pores is not preserved.
Superfamily Monoporelloidea Hincks, 1882
Diagnosis . Colony encrusting (typically multiserial,
sometimes uniserial or pluriserial) or erect (bilaminar-folded
or dichotomously branching with fl exible nodes). Zooids
with opesiules and pores in an extensive cryptocyst, forming
a small opesia coincident with the orifi ce. Gymnocystal rim
Appendices